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Apidae

Apidae is the largest family of bees within the order Hymenoptera and superfamily Apoidea, encompassing approximately 6,000 species distributed across all continents except Antarctica.^[1]^[2] This diverse family includes both social and solitary species, with notable eusocial groups such as honey bees (Apis spp.), bumble bees (Bombus spp.), and stingless bees (Meliponini), as well as solitary genera like carpenter bees (Xylocopa) and long-horned bees (Eucera).^[3] Members of Apidae are characterized by their long tongues adapted for accessing deep floral nectaries, branched body hairs for pollen collection, and varied nesting habits ranging from ground burrows and wood cavities to exposed combs.^[4] The family plays a critical ecological role as primary pollinators for angiosperms, supporting agriculture through species like the managed western honey bee (Apis mellifera), while also exhibiting behaviors such as cleptoparasitism, where some species parasitize the nests of others.^[5] Evolutionarily, Apidae originated in the late Cretaceous around 95 million years ago, with cleptoparasitism emerging multiple times as an ancient adaptation.^[3] Apidae is subdivided into several subfamilies, including Apinae (encompassing honey bees, bumble bees, stingless bees, and many solitary bees), Euglossinae (orchid bees, primarily Neotropical), Xylocopinae (carpenter bees), and Nomadinae (cuckoo bees).^[4] These subfamilies reflect the family's behavioral diversity: eusocial species form complex colonies with division of labor among queens, workers, and drones, producing honey and wax for storage and construction, whereas solitary and parasitic forms lack such structures.^[6] Ecologically, apid bees are vital for biodiversity, with their pollination services contributing to approximately 35% of global crop production, though many species face threats from habitat loss, pesticides, and climate change.^[7]

Characteristics

Morphology

Members of the Apidae family exhibit a typical hymenopteran body plan consisting of a distinct head, thorax, and abdomen. The head is hypognathous, bearing large compound eyes laterally, three ocelli arranged in a triangle on the vertex, and antennae that are geniculate with 12 segments in females and 13 in males. The thorax is robust, comprising the prothorax, mesothorax (with scutum and scutellum), metathorax, and propodeum, with the first abdominal segment fused to form a petiole. Adults possess two pairs of membranous wings: the forewings are larger with three submarginal cells and a closed marginal cell, while the smaller hind wings are attached to the forewings by a row of hook-like hamuli along their anterior margins, facilitating flight efficiency. The body is generally covered in branched hairs, which aid in pollen collection.^[8] Specialized morphological features in Apidae are adapted for nectar feeding and pollen transport. Many species possess a long, flexible proboscis formed by the maxillae and labium, with the glossa (tongue) ranging from about 5 mm in honey bees to over 30 mm in some orchid bees (Euglossini), allowing access to deep floral nectaries.^[9]^[10] Social species, such as those in the genera Apis (honey bees) and Bombus (bumble bees), feature corbiculae—smooth, concave pollen baskets on the outer surfaces of the hind tibiae, fringed with long hairs to hold moistened pollen loads. A key diagnostic trait for female Apidae is the facial fovea, a longitudinal groove paralleling the inner margin of each compound eye, often lined with dense, short hairs. Most females are equipped with a sting apparatus in the abdomen for defense, which is barbed in the subfamily Apinae, causing it to lodge in victims upon deployment.^[8]^[11]^[8] Morphological variation across Apidae subfamilies reflects diverse lifestyles. Members of Xylocopinae, such as carpenter bees (Xylocopa spp.), have robust, large bodies up to 2-3 cm long, with shiny, sparsely haired abdomens and strong mandibles suited for excavating wood nests. In contrast, Nomadinae (cuckoo bees, e.g., Nomada spp.) display slender, wasp-like bodies that are relatively hairless and lack pollen-carrying structures like corbiculae, adaptations for their parasitic habits. Sensory adaptations are consistent across the family, with compound eyes providing panoramic vision for navigation and flower detection, and ocelli sensitive to light intensity for orientation during flight; these structures enable precise foraging and environmental interaction.^[8]^[12]^[13]

Nesting Habits

Apidae exhibit a wide range of nesting strategies, reflecting the family's diverse subfamilies and ecological adaptations, from solitary burrows to complex eusocial colonies. In contrast, species in the subfamily Nomadinae are cleptoparasites that do not build nests but lay eggs in the nests of other bees, relying on host provisions for their offspring.^[11] Many species in the subfamily Apinae construct ground burrows, often in sandy or loose soils, where females excavate vertical tunnels lined with secretions for brood provisioning.^[14] In contrast, Xylocopinae, such as the eastern carpenter bee Xylocopa virginica, nest in wood by excavating unbranched tunnels averaging 15 cm long and 15 mm wide, creating galleries for sequential brood cells partitioned by wood chips and resin.^[15] Bumblebees in the tribe Bombini typically build aerial or subterranean nests in pre-existing cavities like abandoned rodent burrows or under grass tufts, using wax mixed with pollen and plant fibers to form insulating envelopes around brood clusters.^[16] The sociality spectrum in Apidae spans solitary nesters to highly eusocial colonies. Solitary species, such as many Euglossini orchid bees like Euglossa dilemma, construct individual nests in enclosed cavities sealed with resin and plant debris, provisioning cells without assistance from other adults.^[17] Primitively eusocial bumblebees form annual colonies initiated by a single queen, with workers aiding in nest maintenance but lacking rigid caste differentiation, leading to colony decline after one season.^[18] Advanced eusociality is evident in Apini honey bees and Meliponini stingless bees, featuring perennial colonies with distinct castes—queens for reproduction, sterile workers for foraging and care, and drones for mating—supported by overlapping generations. Colony architecture varies markedly across social levels. In Apini, nests consist of parallel wax combs with hexagonal cells for efficient brood rearing and honey storage, optimizing space and material use in cavities like tree hollows.^[19] Meliponini nests feature clustered spherical brood cells and storage pots made from cerumen (wax-resin mixture), often enclosed by resinous batumen walls for protection, with entrance tubes varying by species for controlled access.^[20] Defense mechanisms in social Apidae species emphasize guarding and chemical or physical deterrents. Worker guards at nest entrances antennate intruders to assess threats, releasing alarm pheromones to recruit colony mates for stinging attacks in Apini and Bombini, while Meliponini rely on biting, resin smearing to immobilize predators, and soldier castes for aggressive repulsion.^[21]^[20]

Taxonomy and Classification

Phylogenetic History

The Apidae family originated during the Late Cretaceous period, with fossil evidence indicating their presence around 80 million years ago. The earliest known eusocial apid is Cretotrigona prisca, a stingless bee preserved in New Jersey amber, which represents a worker caste and suggests that advanced sociality had already evolved in the lineage prior to the Cretaceous-Paleogene extinction event approximately 66 million years ago.^[22] This fossil, dating to the Late Cretaceous (approximately 78-92 million years ago), underscores the deep evolutionary roots of eusocial behavior within Apidae, predating the diversification of many modern angiosperms.^[23] Broader bee origins trace back to Western Gondwana around 125 million years ago, but Apidae specifically diversified in the context of emerging flower-bee mutualisms during this era.^[24] Throughout the 19th and 20th centuries, the classification of Apidae underwent significant revisions, including the incorporation of genera from the former families Anthophoridae and Ctenoplectridae into Apidae sensu lato. These mergers, formalized in influential works by Charles D. Michener, reflected growing recognition of phylogenetic affinities based on morphological and early molecular data, expanding Apidae to encompass diverse solitary, social, and parasitic bees. In 2005, Brazilian entomologists Gabriel A.R. Melo and Rodrigo B. Gonçalves proposed a radical unification of all bees (Anthophila) under a single family, Apidae, to align with aculeate Hymenoptera conventions and emphasize monophyly; however, this scheme has not gained widespread acceptance by 2025, as most global classifications retain seven distinct bee families.^[25] Molecular phylogenetic studies have firmly established Apidae as a monophyletic family within the superfamily Apoidea, typically positioned as the sister group to Megachilidae, with robust support from transcriptomic and genomic datasets.^[26] Within Apidae, subfamilies such as Apinae, Xylocopinae, and Nomadinae exhibit confirmed monophyly, with Nomadinae emerging as a derived clade of kleptoparasitic bees that evolved multiple times independently across the family tree.^[27] Key evolutionary adaptations include the transition from solitary nesting to social lifestyles, particularly within Apinae tribes, where eusociality arose once in the corbiculate bees (Apini, Bombini, and Meliponini tribes, including honey bees, bumble bees, and stingless bees) and independently in other lineages such as some Xylocopinae, often linked to corbicular pollen transport and colony defense mechanisms.^[22]^[28] These shifts highlight Apidae's adaptive radiation in response to angiosperm diversification.^[29]

Subfamilies and Tribes

The family Apidae is divided into three subfamilies: Apinae, Nomadinae, and Xylocopinae.^[30] Apinae is the largest subfamily, encompassing a diverse array of eusocial and solitary bees, with key tribes including Apini, Bombini, Meliponini, as well as other tribes like Euglossini (orchid bees) and Anthophorini (digger bees). The tribe Apini comprises the honey bees of the genus Apis, which includes 9 extant species such as Apis mellifera, the western honey bee widely managed for pollination and honey production.^[31] Bombini consists of bumblebees in the genus Bombus, totaling approximately 250 species that serve as robust pollinators in temperate and boreal regions due to their ability to buzz-pollinate flowers.^[24] Meliponini includes stingless bees, with around 600 described species primarily distributed in tropical regions, where they produce honey and exhibit advanced eusocial behaviors in cavity nests.^[32] Nomadinae, the subfamily of cuckoo bees, contains over 1,500 species that are obligate kleptoparasites, invading the nests of other bees to lay eggs and lacking specialized pollen-collecting structures like scopae; prominent tribes include Nomadini, with genera such as Nomada.^[33] Xylocopinae features relatively large-bodied bees, with tribes such as Xylocopini, which includes carpenter bees that nest in wood by excavating galleries, and Allodapini, comprising small communal species that exhibit subsocial behaviors in stems or twigs.^[30]

Life Cycle and Reproduction

Developmental Stages

Members of the Apidae family undergo complete metamorphosis, progressing through four distinct developmental stages: egg, larva, pupa, and adult. This holometabolous life cycle is characteristic of all bees in the family, enabling adaptation to diverse ecological niches, from solitary to highly eusocial species. The duration and specifics of each stage vary by species, caste (in social groups), and environmental factors such as temperature, but the process ensures the transformation from a dependent juvenile to a fully functional adult pollinator.^[8] The egg stage begins when a female bee—typically the queen in social species like honey bees (Apis mellifera) or bumble bees (Bombus spp.)—lays tiny, elongated, white, cylindrical eggs, each about 1-2 mm long, into individual cells within the nest. These eggs are often deposited on or near a provision of food, such as a pollen-nectar mixture. In social Apidae, the incubation period lasts 3-5 days; for example, in A. mellifera, eggs hatch after approximately 3 days, while in B. impatiens, hatching occurs in about 4 days. Solitary Apidae females lay a single egg per cell before sealing it.^[8]^[34] Hatching reveals the larval stage, where legless, C-shaped, pale grubs emerge, initially measuring less than 2 mm but growing rapidly through feeding and molting. Larvae in eusocial species, such as Apis and Bombus, receive progressive provisioning via trophallaxis—mouth-to-mouth transfer of glandular secretions, nectar, and pollen from adults—allowing open brood care in shared chambers or cells. In contrast, many solitary Apidae employ mass or closed provisioning, where the female pre-loads the cell with a pollen-nectar paste (beebread) before laying the egg and sealing it. Larvae typically undergo 5-6 instars, molting their exoskeleton each time to accommodate growth; durations vary, lasting 5-6 days in A. mellifera workers or up to 10-14 days in B. impatiens. In solitary species like carpenter bees, larvae may enter diapause during winter, extending the cycle seasonally.^[8]^[35]^[34] The pupal stage follows when the mature larva spins a silken cocoon within the cell, forming a non-feeding chrysalis where internal reorganization occurs, developing compound eyes, wings, legs, and genitalia. This enclosed phase protects the developing adult and lasts 7-14 days, influenced by species and temperature; for instance, it takes about 11-12 days in A. mellifera workers and roughly 2 weeks in B. impatiens. The pupa gradually darkens and assumes the adult form, with no further nutrient intake required after cell capping by workers in social species.^[8]^[34] Adult emergence, or eclosion, completes the cycle as the fully formed bee chews through the cell cap or partition using its mandibles, typically after 21 days total in A. mellifera workers or 4-5 weeks in B. impatiens. Sexual dimorphism becomes apparent at this point: males (drones) lack a functional sting apparatus (derived from an ovipositor present in females) and often exhibit size differences, such as being larger than workers in Apis but similar in Bombus, while females include sterile workers and fertile queens with stings for defense. Newly emerged adults are pale and soft but harden quickly, ready to assume roles in foraging, nursing, or reproduction depending on caste and species.^[8]^[34]^[35]

Mating Behaviors

Mating systems in Apidae vary widely across subfamilies, reflecting adaptations to sociality, habitat, and ecology. In eusocial species like honey bees (Apis mellifera in the tribe Apini), queens are polyandrous, mating with multiple drones (typically 10–20) during a single nuptial flight early in adulthood, after which they store sufficient sperm for lifelong egg production without further mating.^[36] This strategy ensures genetic diversity in the colony while minimizing exposure to predation during flights. In contrast, bumble bee queens (Bombus spp., also in Apini) are predominantly monandrous, mating once with a single male, though polyandry occurs in some species such as Bombus hypnorum, where queens may mate with up to three males, potentially enhancing colony fitness through increased patriline diversity.^[37] Among the orchid bees (Euglossini), mating follows a lekking system where males aggregate at display sites, such as treefalls or vertical branches in neotropical forests, to expose volatile perfume blends collected from orchids and other sources; females visit these leks to select and mate with males based on fragrance quality and display vigor, without resource provisioning by males.^[38] Queen production in eusocial Apidae involves specialized rearing processes. In honey bees (tribe Apini), workers select young larvae and feed them exclusively with royal jelly—a protein-rich glandular secretion—in enlarged queen cells, promoting development into larger, reproductively capable individuals.^[39] In bumble bees (tribe Bombini), the first-generation larvae that become queens receive superior amounts of pollen and nectar provisions compared to workers, without the use of royal jelly. In stingless bees (Meliponini), queens develop in dedicated cells provisioned with nutrient-rich larval food, often from larger or genetically predisposed larvae.^[40] Upon emergence, virgin queens undertake nuptial flights to mate: in honey bees, these occur shortly after eclosion, with the queen flying to drone congregation areas up to several kilometers from the nest; similarly, bumble bee queens mate on the ground or low vegetation post-hibernation. This brief mating phase is critical, as mated queens return to initiate or join colonies, while unmated ones perish. Colony reproduction strategies differ markedly among eusocial groups. In Apini such as honey bees, reproduction occurs via swarming, where the prime queen departs with a large swarm of workers (often 50–70% of the colony) to establish a new nest, leaving behind developing queens to head the parent colony after conflict resolution.^[41] Stingless bees (Meliponini) employ budding, a gradual fission process where a portion of workers, including the queen or a new queen, slowly relocate to a nearby site, often within meters, allowing the mother colony to retain most resources and minimize risk. Solitary Apidae species, such as many carpenter bees (Xylocopinae), produce multiple broods annually; females construct sequential nests, laying eggs in provisions and overwintering to repeat the cycle, with one to several generations per year depending on climate and resources.^[42] Parasitic strategies in the subfamily Nomadinae represent an alternative reproductive mode, where females act as cleptoparasites by invading host nests—often of ground-nesting Andrenidae or other Apidae—to lay eggs directly into provisioned cells without establishing their own nests or engaging in host-specific mating cues beyond normal insemination by males.^[43] The parasitic larvae consume the host's provisions upon hatching, developing rapidly to maturity without worker caste formation, thus bypassing typical colony investment.

Ecology

Global Distribution

The family Apidae has a cosmopolitan distribution, present across all continents except Antarctica, though it is absent from extreme polar regions in its native ranges, with introduced species extending into some colder areas. The highest species diversity occurs in tropical and subtropical zones, reflecting the family's evolutionary origins and adaptation to warm climates.^[44] However, climate change is driving shifts in Apidae distributions as of 2025, with projections indicating that about 65% of bee species may experience reduced suitable ranges, particularly in Africa and Europe, while some temperate species expand poleward.^[45] Subfamily distributions within Apidae show distinct biogeographic patterns. The Apinae, the largest subfamily, predominates in both temperate and tropical regions worldwide; for example, the tribe Bombini (bumblebees) is primarily native to the Holarctic realm, while Meliponini (stingless bees) exhibit a pantropical range with the greatest diversity in the Neotropics. The Xylocopinae are largely pantropical, occurring mainly in warm, humid environments across Africa, Asia, Australia, and the Americas. Nomadinae, being cleptoparasitic, have distributions that closely track those of their host bees, resulting in a cosmopolitan but patchy presence aligned with Apinae and other host-rich areas.^[46]^[47]^[48]^[49]^[50] Human-mediated introductions have significantly altered Apidae distributions, most notably with Apis mellifera, the western honey bee, which is native to Europe, the Middle East, and Africa but has been transported globally to every continent except Antarctica, establishing feral populations in regions like the Americas, Australia, and Asia. These introductions often impact native ecosystems by competing with local pollinators for resources and altering plant-pollinator dynamics.^[51]^[52]^[53] Apidae species occupy diverse habitats, including forests, grasslands, and urban areas, demonstrating broad environmental adaptability. They range altitudinally from sea level to high elevations, with some species, such as Andean bumblebees (Bombus spp.), inhabiting altitudes up to 4,750 meters in the Andean mountains.^[54]^[55]

Environmental Interactions

Members of the Apidae family play a pivotal role in ecosystems through their pollination services, which facilitate plant reproduction and maintain biodiversity. However, climate change is disrupting these interactions through phenological mismatches, where bees and flowering plants bloom out of sync, potentially reducing pollination efficiency.^[56] Bumblebees in the tribe Bombini are renowned for buzz pollination, a specialized mechanism where they grip flowers of solanaceous plants, such as tomatoes and blueberries, and generate rapid thoracic vibrations—known as sonication—to dislodge pollen from poricidal anthers.^[57]^[58] This vibration, produced by the bee's indirect flight muscles at frequencies around 300-400 Hz, efficiently extracts pollen that would otherwise be inaccessible, enhancing pollen transfer to stigmas during the process.^[59] In contrast, orchid bees of the tribe Euglossini exhibit a unique specialization, where males actively collect volatile fragrances from orchid flowers using their forelegs and midlegs, inadvertently transferring pollinia between plants.^[60] These scents, often complex mixtures of terpenoids and aromatics, serve as attractants that drive male visitation, ensuring pollination for over 700 Neotropical orchid species while the bees store the compounds in specialized hind tibial organs for later use in mating displays.^[61]^[62] Apidae species are integral to trophic interactions, serving as prey and hosts in food webs. Adult bees, including honeybees and bumblebees, are targeted by predators such as birds (e.g., bee-eaters and shrikes) and spiders (e.g., crab spiders like Misumena vatia), which ambush foraging individuals at flowers, reducing bee populations and influencing foraging behaviors like increased vigilance or shorter visit durations.^[63]^[64] Parasitic relationships are exemplified by the mite Varroa destructor, an ectoparasite of Apis mellifera, which attaches to immature and adult bees to feed on hemolymph, weakening hosts, transmitting viruses, and causing colony collapse if unmanaged.^[65] Within the family, kleptoparasitism occurs via the subfamily Nomadinae, or cuckoo bees, which infiltrate nests of ground-nesting hosts like Andrena or Halictus species, laying eggs that hatch into larvae consuming the host's provisions without contributing to nest construction.^[66]^[67] Symbiotic associations further underscore Apidae's ecological integration. Gut microbiomes in corbiculate bees, such as those dominated by Lactobacillus and Bifidobacterium in honeybees and bumblebees, aid digestion by fermenting complex carbohydrates from pollen and nectar, producing short-chain fatty acids that enhance nutrient absorption and provide pathogen resistance.^[68]^[69] Plant-bee coevolution has shaped morphological adaptations, as seen in long-proboscid bees like those in the genus Xylocopa or certain Anthophora species, whose extended mouthparts match the deep corolla tubes of specialized flowers (e.g., in Lamiaceae or Orchidaceae), ensuring exclusive access to nectar and precise pollen deposition.^[70]^[71] These interactions yield broader ecosystem impacts, particularly in enhancing plant reproduction and soil health. Pollination by Apidae boosts seed set in diverse flora; for instance, bee visitation increases seed production by up to 30-60% in crops like sunflowers and wild plants, promoting genetic diversity and habitat stability.^[72]^[73] Ground-nesting species, including many solitary Apidae like Nomia and Melissodes, contribute to soil aeration by excavating burrows that improve porosity, water infiltration, and microbial activity, thereby supporting nutrient cycling and reducing erosion in grasslands and agricultural margins.^[74]

Diversity and Examples

Species Counts

The family Apidae includes approximately 6,000 described species as of 2025, though this figure is likely an underestimate given the abundance of undescribed taxa in tropical regions.^[75] Recent taxonomic efforts, including DNA barcoding initiatives, continue to reveal new species, contributing to ongoing revisions in species counts, with global bee diversity estimates suggesting further increases.^[76] Species richness within Apidae is unevenly distributed across its subfamilies, with Apinae accounting for approximately 3,500 species, Nomadinae around 1,500, and Xylocopinae about 1,200.^[77] These breakdowns highlight Apinae's dominance, encompassing diverse groups such as honey bees, bumble bees, and stingless bees, while Nomadinae consists primarily of cleptoparasitic cuckoo bees and Xylocopinae features wood-nesting carpenter bees. Diversity hotspots for Apidae are concentrated in the Neotropics, particularly for the tribe Meliponini within Apinae, where hundreds of stingless bee species thrive in rainforests.^[78] In contrast, the Holarctic region serves as a key center for Bombini (bumble bees), with significant species accumulation in temperate zones.^[2] Advances in DNA barcoding have accelerated discoveries in these areas, uncovering cryptic diversity and refining phylogenetic relationships.^[66] Endemism patterns in Apidae are pronounced on islands, such as in the Neotropics where many Meliponini species are endemic to specific archipelagos. Such isolated populations contribute uniquely to global counts but face pressures from habitat loss and invasive species, potentially altering future tallies of extant diversity.^[79]

Notable Groups

Within the family Apidae, the tribe Apini is exemplified by Apis mellifera, the Western honey bee, which is widely managed across continents for its eusocial colony structure and pollination services. Worker bees in A. mellifera colonies forage extensively, contributing to honey production, with each worker yielding approximately 1/12 teaspoon (about 500 mg) of honey over her 6-week summer lifespan.^[80]^[81] The tribe Bombini features Bombus terrestris, a common bumblebee known for its robust, hairy body adapted to temperate climates. Queens of B. terrestris hibernate overwinter in soil cavities, emerging in spring to establish colonies that support buzz pollination in various crops.^[82] In the tribe Meliponini, Melipona beecheii represents stingless bees native to Mesoamerica, where it has been cultivated since pre-Columbian times by Mayan communities for its honey. Lacking stings, M. beecheii colonies defend their nests using resin collected from plants, which workers apply in bites to deter intruders.^[83]^[84] The tribe Xylocopini includes Xylocopa virginica, the eastern carpenter bee, a large solitary species that excavates tunnels in soft, weathered wood for nesting. Females of X. virginica provision cells with pollen and nectar for their larvae, exhibiting primitive social tendencies in some aggregations.^[80] Euglossini is highlighted by Euglossa dilemma, an orchid bee characterized by its striking metallic blue coloration. Males of E. dilemma actively collect volatile fragrances from orchids and other sources, storing them in hind-leg cavities to create perfumes that attract females during lekking displays. Nomadini encompasses Nomada spp., kleptoparasitic cuckoo bees with slender, wasp-like bodies and reduced wing venation. Females of Nomada species infiltrate nests of ground-nesting Andrena bees, laying eggs that hatch into larvae which consume the host's provisions.^[85]

Human Significance

Economic Roles

Apidae species, particularly honey bees (Apis spp.) and bumble bees (Bombus spp.), play a pivotal role in the global pollination economy by supporting the production of numerous crops. In the United States, honey bees pollinate crops valued at nearly $20 billion annually, including key commodities such as almonds and apples that rely heavily on managed bee colonies for consistent yields.^[86] Bumble bees contribute significantly to the pollination of crops like tomatoes through buzz pollination, a specialized vibration technique that enhances fruit set in greenhouse production, supporting an industry with global market value exceeding $200 billion as of 2024.^[87] Overall, insect pollination services, dominated by Apidae, generate an estimated over $800 billion in annual economic value worldwide for agricultural output as of 2025.^[88] Beekeeping with Apidae yields a range of valuable products, with honey being the primary commodity derived from species like the Western honey bee (Apis mellifera) and stingless bees (e.g., Melipona spp.). Global honey production reached about 1.9 million metric tons in 2023, driven largely by commercial apiculture in regions such as Asia and Europe. Other products include beeswax, used in cosmetics and candles; propolis, valued for its antimicrobial properties in pharmaceuticals; and royal jelly, harvested for its nutritional content in health supplements. These outputs form a multibillion-dollar industry, with the global honey market alone valued at around $9.2 billion in 2024. Commercial rearing of Apidae focuses on select managed species to meet agricultural and product demands. Apis mellifera is the cornerstone of worldwide beekeeping operations, with millions of hives maintained for pollination and honey production. In North America, Bombus impatiens is commercially reared for greenhouse pollination, particularly for crops requiring buzz pollination like tomatoes and peppers. In tropical regions, stingless bee farming—known as meliponiculture—involves species such as Melipona beecheii, providing honey and pollination services while supporting local economies in Mesoamerica. Beyond direct economic uses, Apidae hold cultural significance in human societies, symbolizing royalty and divinity in ancient traditions. In ancient Egypt, bees were sacred emblems associated with the pharaohs and deities like Ra, representing the soul's journey and appearing in hieroglyphs as early as 3500 BCE. Additionally, bee venom from Apidae species is employed in apitherapy, a medicinal practice using controlled stings or injections to alleviate conditions such as arthritis and inflammation, with ongoing research exploring its anti-inflammatory and analgesic effects.

Conservation Challenges

Apidae populations face multiple anthropogenic and environmental threats that contribute to widespread declines. Habitat loss due to deforestation and urbanization is a primary driver, fragmenting foraging areas and nesting sites essential for species like bumblebees and solitary bees within the family.^[89] Pesticides impair foraging behavior and reproductive success by contaminating pollen and nectar, leading to sublethal effects on navigation and colony health, with neonicotinoids posing particular risks.^[90]^[91] Climate change exacerbates these issues by shifting plant bloom times out of sync with bee life cycles, reducing food availability and altering suitable habitats for thermoregulation-sensitive species.^[92] Diseases such as Nosema ceranae infection weaken honey bee (Apis spp.) colonies by disrupting gut function and shortening worker lifespan, often interacting with stressors like poor nutrition to cause colony collapse.^[93] In 2024-2025, managed honey bee colony losses in the US reached a record 55.6%, threatening pollination services critical to $17 billion in agricultural production.^[94] Invasive species, including Africanized honey bees (Apis mellifera scutellata hybrids), compete aggressively for resources and hybridize with native populations, reducing genetic diversity in Neotropical Apidae communities.^[95] Certain Apidae groups are particularly vulnerable, highlighting the uneven impact of these threats. In North America, approximately 28% of bumble bee (Bombus spp.) species are considered threatened with extinction, driven by habitat degradation and pathogen spillover from managed bees.^[96] Stingless bees (Meliponini tribe) in tropical regions face heightened risks from deforestation and agricultural intensification, which destroy resin sources and nesting cavities in dry forests and rainforests.^[97] These declines underscore the broader pollinator crisis, with 172 wild bee species in Europe now assessed as threatened as of 2025, many belonging to Apidae subfamilies.^[98] Conservation efforts target these threats through a combination of policy, habitat restoration, and research. Protected areas and pollinator gardens enhance floral diversity and nesting opportunities, supporting resilient populations of native Apidae.^[99] The European Union has implemented strict regulations, banning outdoor use of three neonicotinoids (imidacloprid, clothianidin, thiamethoxam) since 2018 to mitigate pesticide exposure, with further court rulings in 2023 prohibiting temporary exemptions.^[91] Breeding programs for disease resistance, including selection against Nosema-susceptible strains in honey bees, aim to bolster colony survival, while nutritional supplementation reduces infection severity.^[100] The IUCN Red List has evaluated over 200 bee species globally, including numerous Apidae, informing targeted protections like those for North American Bombus.^[101] Global initiatives, such as the Pollinator Partnership's habitat certification and education programs, promote widespread adoption of bee-friendly practices.^[99] Recent IPBES assessments, including the 2025 Transformative Change report, emphasize urgent interventions to reverse pollinator declines, projecting continued biodiversity loss without scaled-up action.^[102]

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Nov 26, 2020 · We synthesise the literature on the nesting-habitat associations of ground-nesting bees, focussing on soil characteristics.Bee Life History · Nest Construction · Abiotic Factors
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Eastern Carpenter Bee (Hymenoptera: Apidae): Nest structure, nest ...
Mean values of X. virginica nest measurements (tunnel length 15.4 ± 1.2 cm, width 15.0 ± 0.5 mm, and cell length 17.7 ± 0.3 mm) were similar to those reported ...
[13]
Bumble bee nesting 101
Bumble bees often nest in abandoned rodent dens underground or under tufts of grass. Rodents create spacious shelters and fill them with insulating materials, ...Missing: habits | Show results with:habits
[14]
Green Orchid Bee Euglossa dilemma Friese (Insecta: Hymenoptera
Nests are located in any sort of enclosed cavity; entrances are often sealed off with resin and plant debris (Figure 4a and 4c). Mature nests can contain up to ...<|control11|><|separator|>
[15]
Bumble Bees | USU
Nesting Habits. social; colony dies off every fall; fertilized queens overwinter, usually underground; nest in old rodent burrows, holes, grass clumps, etc.
[16]
Imperfect comb construction reveals the architectural abilities of ...
Aug 12, 2021 · Honeycomb is hailed as the pinnacle of biological architecture for its ability to maximize storage area while minimizing building material (1–4) ...
[17]
Stingless bee nesting biology* - Apidologie
Inability to freely swarm and single mating may all increase nesting specificity, competition, symbiosis and cleptobiosis in communities, while disease is rare.
[18]
The defensive response of the honeybee Apis mellifera
Nov 15, 2016 · Intra-specific defence. When another bee lands at the hive entrance, guards quickly approach and antennate it in order to check whether it is a ...
[19]
The Antiquity and Evolutionary History of Social Behavior in Bees
We conclude that eusociality evolved once in the common ancestor of the corbiculate Apidae, advanced eusociality evolved independently in the honey and ...
[20]
Family-Group Names for Bees (Hymenoptera: Apoidea) - BioOne
Perhaps the longest used name for the bees has been Anthophila as proposed by Latreille (1804: originally in the plural as Anthophili, but subsequently emended: ...
[21]
The evolutionary history of bees in time and space - ScienceDirect
Aug 21, 2023 · We present a novel analysis of bee biogeography using extensive new genomic and fossil data to demonstrate that bees originated in Western Gondwana (Africa and ...
[22]
https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0021086
[23]
Phylogenomic analysis of Apoidea sheds new light on the sister ...
May 18, 2018 · Our phylogenetic result, based on analyzing a combined enriched and transcriptomic dataset, allowed to trace the evolutionary origin of bees and ...
[24]
Comprehensive phylogeny of apid bees reveals the evolutionary ...
We recover a monophyletic Xylocopinae and Nomadinae and find that Osirini, Protepeolini, and Isepeolini should not be placed within Nomadinae. Previous ...Results And Discussion · Phylogenetic Relationships · Materials And Methods
[25]
Phylogeny of the carpenter bees (Apidae: Xylocopinae) highlights ...
Aug 30, 2023 · Our results indicate solitary origins for the Xylocopinae with multiple transitions to sociality across the tree and subsequent reversals to solitary life.
[26]
Family APIDAE - Australian Faunal Directory
Apr 30, 2025 · The family is divided into three subfamilies, the Apinae, the Nomadinae and the Xylocopinae. The largest subfamily, Apinae, is further divided ...
[27]
Apis Linnaeus, 1758 - GBIF
Only eight surviving species of honey bee are recognized, with a total of 43 subspecies, though historically 7 to 11 species are recognized. Honey bees ...Missing: count | Show results with:count
[28]
Stingless bee classification and biology (Hymenoptera, Apidae): a ...
In addition to the systematic review, the biology of stingless bees is summarized with an emphasis on aspects related to their nesting biology and architecture.
[29]
Apidae) with integration of UCE and DNA barcode sequence data
May 17, 2025 · Among the many independent clades of parasitic bees, by far the most species is the subfamily Nomadinae, containing over 1,500 species divided ...
[30]
EENY-777/IN1348: Himalayan Giant Honey Bee, Cliff Honey Bee ...
Feb 25, 2022 · The life of a honey bee starts when an egg laid by the queen in a wax cell. The larva will eclose (emerge) from the egg after three days. Larva.
[31]
[PDF] Bombus impatiens, Common Eastern Bumble Bee - LSU AgCenter
bees emerge from the cocoons. The entire process of growth from egg to larva, pupa and adult takes around four to five weeks, depending on weather ...
[32]
[PDF] Wild Bees of Eastern North America - Rehan Lab
stages: egg, larva, pupa, and adult. Most solitary mother bees build a single cell, provision it with a ball made of pollen and nectar, lay a single egg on it,.
[33]
Observation of the Mating Behavior of Honey Bee (Apis mellifera L ...
Jul 1, 2014 · Honey bee (genus Apis/Apis mellifera) queens are polyandrous and mate only at one period in their life [1,2,3,4,5]. Usually within 1 week after ...
[34]
Dynamics of multiple-mating in the bumble bee Bombus hypnorum
The mating systems of social insects range from monandry, through low levels of polyandry to extremely high levels of polyandry. While the distribution of ...
[35]
The behaviour of male orchid bees (Apidae, Hymenoptera, Insecta ...
The relationships between male territories, female foraging ranges, and collection of aromatic material by males are discussed. Recommended articles ...Missing: paper | Show results with:paper
[36]
Male mating behaviour and mating systems of bees: an overview1
Jun 1, 2005 · To maximise their reproductive success, males will therefore be selected to maximise their access to recep- tive (immediate post-emergence) ...
[37]
[PDF] Reproduction in eusocial bees - (Apidae: Apini, Meliponini) - DSpace
Swarming involves the departure of a cohort of workers with a queen that leaves the original colony for a new nest site. In honeybees, the old queen leaves the ...
[38]
Garden Allies: Bees II (Solitary to Social) - Pacific Horticulture
Apidae have the broadest range of social behaviors of any bee family, exhibiting the full gamut from solitary to highly eusocial.
[39]
[PDF] Nomadinae - Squarespace
Oct 16, 2021 · hosts of nomadine parasites. Similarly, these techniques shed light on the most likely behavioral strategies used by the first brood parasitic.
[40]
Biogeography: The origin and spread of bee lineages - ScienceDirect
Aug 21, 2023 · A new study that combines phylogenetics with fossil data reconstructs the origin and diversification of bees across geological time and space.
[41]
The bee tree of life: a supermatrix approach to apoid phylogeny and ...
Jul 3, 2013 · Biogeography. The current distributions of bee genera were assigned to one or more of seven broad geographic regions: Africa, Eastern Palearctic ...
[42]
Biogeography and Diversification of Bumblebees (Hymenoptera
According to this model, the most likely biogeographic distribution of the most recent common ancestor (MRCA) of all bumblebees was in the Eurasian (Palearctic ...
[43]
Phylogenomic insights into the worldwide evolutionary relationships ...
Phylogenomic data are generated for all genera of stingless bees worldwide. · We provide a reliable phylogenetic basis to investigate the evolution of the group.<|control11|><|separator|>
[44]
Phylogenomics and biogeography of the small carpenter bees (Apidae
The apid subfamilies Eucerinae, Apinae, Nomadinae, and Anthophorinae each independently appear as progressively more distant sister groups to Xylocopinae. The ...2. Materials And Methods · 3. Results · 4. Discussion<|separator|>
[45]
Host‐Cleptoparasite Biogeographical Congruence Through Time
Feb 6, 2025 · 2019). Within Apidae, the subfamily Nomadinae is notable for being the most diverse (about 1200 species) and ancient (about 100 Ma) lineage of ...
[46]
European Honey Bee Apis mellifera Linnaeus and subspecies ...
The western honey bee, Apis mellifera Linnaeus, naturally occurs in Europe, the Middle East, and Africa. This species has been subdivided into at least 20 ...
[47]
Apis mellifera (European honeybee) | CABI Compendium
Nov 10, 2022 · Currently, managed colonies as well as naturalized feral colonies of A. mellifera occur in every continent except for Antarctica. Apis mellifera ...
[48]
Density of invasive western honey bee (Apis mellifera) colonies in ...
Mar 4, 2022 · Feral Apis mellifera colonies are widespread globally and cause ecological impacts as pollinators and competitors for food and nesting ...
[49]
Nesting Biology of Xylocopa xinjiangensis (Hymenoptera: Apidae
Jul 5, 2018 · The species within Xylocopa are distributed throughout the world except for Antarctica, mainly in tropical and subtropical regions, with a few ...
[50]
High thermal tolerance in high‐elevation species and laboratory ...
Dec 4, 2022 · Among these species, B. funebris reaches the highest elevations in the Andes, as it has been recorded at 4750 m in Colombia (Gonzalez & Engel, ...
[51]
Buzz-Pollinated Crops: A Global Review and Meta-analysis of the ...
Feb 22, 2021 · Buzz-pollinated plants require visitation from vibration producing bee species to elicit full pollen release. Several important food crops are ...
[52]
Buzz pollination: studying bee vibrations on flowers - Vallejo‐Marín
Dec 26, 2018 · Buzz pollination is a type of pollination in which bees use vibrations to remove and collect pollen from flowers incidentally fertilising them.Missing: Bombini solanaceous
[53]
(PDF) Buzzing Bees (Hymenoptera: Apidae, Halictidae) on Solanum ...
Aug 6, 2025 · The anther cone needs a vibratory motion (buzz pollination or sonication) to expel the pollen onto the female flower parts (De Luca & Vallejo- ...
[54]
Function of environment-derived male perfumes in orchid bees
May 22, 2023 · Here, we demonstrate that the possession of perfume increases male mating success and paternity in Euglossa dilemma, a species of orchid bees ...
[55]
Phylogeny and biology of neotropical orchid bees (Euglossini)
Males are especially important in the pollination of some orchids, from which they collect aromatic fragrances thought to play a role in territorial display and ...Missing: scents | Show results with:scents
[56]
Composition of Orchid Scents Attracting Euglossine Bees
Aug 5, 2025 · In the species that are exclusively pollinated by fragrance‐collecting male euglossine bees, the fragrances are primary attractants that serve ...<|separator|>
[57]
effects of predation risk from crab spiders on bee foraging behavior
In this paper, we explore the effects of crab spiders (Araneae: Thomisidae) on the behavior of 2 species of bee (Hymenoptera: Apidae) foraging for nectar and ...
[58]
Impact of prédation upon honey bees (Hymenoptera, Apidae), by ...
Estimates of prédation on the honey bee, Apis mellifera L., by a guild of orb-weaving spiders were made in two different habitats in central Europe.
[59]
Varroa mite - Carl Hayden Bee Research Center - USDA ARS
Apr 5, 2023 · The Varroa mite (Varroa destructor) is an ectoparasite of the honey bee Apis mellifera L. Varroa is the most serious pest of honey bees ...
[60]
Harrison's rule corroborated for the body size of cleptoparasitic ...
Jun 29, 2022 · Nomadinae, with nearly 1,300 described species globally, is the largest lineage where all members are obligatory brood parasites27.
[61]
Cuckoo bees - Florida Wildflower Foundation
Cuckoo bees are often mistaken for wasps because their body shape resembles a wasp, and they are nearly hairless. They also lack the pollen baskets.Missing: morphology slender<|control11|><|separator|>
[62]
Symbiosis of Carpenter Bees with Uncharacterized Lactic Acid ...
Jun 22, 2023 · Gut microbiomes of the family Apidae have been extensively examined in eusocial bees such as honey bees and bumble bees, and are composed of bee ...
[63]
The honey bee gut microbiota: strategies for study and ...
Jan 16, 2019 · This review discusses methods and pipelines used in the study of the gut microbiota of Ap. mellifera and closely related species for four main purposes.
[64]
Drinking with a very long proboscis: Functional morphology of orchid ...
Bee mouthparts form a proboscis, which is often elongated to access nectar from long-spurred flowers. Euglossini or orchid bees are a fascinating group of ...
[65]
Fine-tuned Bee-Flower Coevolutionary State Hidden within Multiple ...
Feb 5, 2014 · These results demonstrate a fine one-to-one morphological match between the bee's proboscis and the corolla tube. Such morphological ...
[66]
Pollinators significantly enhance seed set, yields and chemical ...
Mar 1, 2025 · This study highlights the positive impact of floral insect visitors on sunflower seed set, seed mass, crop yield, seed oil content, and fatty acid composition.
[67]
Bee-mediated pollination enhances fruit set and seed yield in ...
Nov 1, 2025 · Bee pollination boosts agricultural productivity, food security, and livelihoods by increased fruit, seed, and oil yields while reducing ...
[68]
Regard and protect ground‐nesting pollinators as part of soil ...
Feb 9, 2022 · Ground-nesting pollinators change soil texture directly by digging cavities, but more importantly by their indirect contribution to soil quality and functions.
[69]
Apidae – López-Uribe Lab
Jun 29, 2020 · Three submarginal cells on the forewing · Long tongue length · Small to very large size · Wide range of colors and patterns · Many species are ...Missing: diagnostic | Show results with:diagnostic
[70]
How many bee species are there? A quantitative global estimate
Jun 27, 2025 · Globally, we estimated 24,705–26,164 bee species, an 18–25% increase, representing at least 32–45 years of taxonomic research. Particularly high ...
[71]
https://pmc.ncbi.nlm.nih.gov/articles/PMC3913927/
[72]
Stingless bee classification and biology (Hymenoptera, Apidae)
Jul 27, 2023 · The closest relatives of Meliponini are the species of the extinct tribe Melikertini ( Engel 1998 , 2001a , 2001b ; Schultz et al. 2001 ), a ...
[73]
Bee Diversity — Museum of the Earth
There are seven families of bees. Each family has its own set of shared traits and a common evolutionary history.Missing: multiple | Show results with:multiple
[74]
Subfamily Xylocopinae - Carpenter and Small carpenter Bees
33 spp. in 2 genera of 2 tribes in our area, >1000 spp. in 16 genera of 4 tribes total(1) Range worldwide; our tribes are both cosmopolitan.
[75]
Apidae - GBIF
This is a test site. The production site with full data is available at GBIF.org. Continue to test site. Apidae. Dataset; GBIF Backbone Taxonomy: Rank ...Missing: URL | Show results with:URL
[76]
https://www.researchgate.net/publication/392856655_How_many_bee_species_are_there_A_quantitative_global_estimate
[77]
Ambler Arboretum is Abuzz About Bees with the Return of ...
Jan 9, 2024 · An average worker bee will only make 1/12 of a teaspoon of honey in its lifetime. One honeybee colony can produce 60 to 100 pounds of honey per ...
[78]
Bombus terrestris (bumble bee) | CABI Compendium
The first bumble bees to be seen in spring are the queens – the queen is the only bumble bee to hibernate through the winter. ... bumble bees for greenhouse crop ...
[79]
(PDF) Extinction of Melipona beecheii and traditional beekeeping in ...
Aug 6, 2025 · Beekeepers using this bee, from the Mayan zone in Quintana Roo state, Mexico, testify to a 93% decrease in hives during the past one-quarter century.
[80]
Resin Use by Stingless Bees: A Review - PMC - PubMed Central
Resin use is vital to many aspects of stingless bee colony function. Stingless bees use resin to build essential nest structures, repel predators, and kill ...
[81]
Spring Bees: Who Are They and Where Do They Live?
Jun 4, 2023 · ... nests, eat the egg of the host, and lay their own egg. Common cuckoo bees include bees in the genus Nomada, which may be confused with wasps ...
[82]
Rusty Patched Bumble Bee - Xerces Society
Major threats that alter landscapes and habitat required by bumble bees include agricultural and urban development. Livestock grazing also may pose a threat to ...
[83]
Threats to Pollinators | Pollinator.org
Threats to Pollinators · Threats to Pollinators And why they need our help · Habitat Loss · Climate Change · Misuse of Pesticides · Parasites and Pathogens · Non- ...
[84]
More time to fly: with a warming climate the Western honey bee ...
Warming temperatures (ignoring all other factors), may lead to more flight time for honey bees; may affect plant bloom times and species distributions; and may ...
[85]
Nosema Disease - Bee Research Laboratory - USDA ARS
Aug 12, 2016 · Nosema disease is widespread and causes serious damage to adult honey bees thus reducing the life span of individual bees and weakening or killing colonies.Missing: conservation | Show results with:conservation
[86]
Africanized Honeybee | National Invasive Species Information Center
African honey bee, Africanized honey bee, or killer bee, Apis mellifera scutellata Lepeletier (Hymenoptera: Apidae). In: J.L. Capinera (Ed.), Encyclopedia of ...
[87]
Factors Contributing to Bee Decline | USU
In North America, 26% of bumble bee species (Bombus spp.) are considered threatened (i.e., likely to be listed as endangered under the ESA), with widespread ...
[88]
Full article: Stingless bees in tropical dry forests: global context and ...
This paper aims to provide a global perspective of the relationship between the breeding of stingless bees and the conservation of tropical dry forests.
[89]
[PDF] European Red List of Bees - IUCN Portal
family of bees is the Apidae (561 species), which includes the Honey bee and ... of life (Apidae, Halictidae and Megachilidae). Cuckoo bees can be very ...<|control11|><|separator|>
[90]
Pollinator Partnership
A non-profit 501(c)3 organization-- the largest in the world dedicated exclusively to the protection and promotion of pollinators and their ecosystems. OUR ...Planting Guides · Pollinator Week · Posters · About Us
[91]
Neonicotinoids - Food Safety - European Commission
In 2013, the Commission severely restricted the use of plant protection products and treated seeds containing three of these neonicotinoids (clothianidin, ...
[92]
What's new with Nosema treatment? - UF/IFAS Entomology ... - Blogs
Sep 23, 2019 · Nosema is treatable. It is recommended that beekeepers ensure that their colonies have access to adequate nutrition. Adequate feeding may decrease the negative ...
[93]
[PDF] IUCN Assessments for North American Bombus spp. - Xerces Society
We assigned each species to a preliminary IUCN Red List Category (IUCN 2012) by looking at ... bee (Apidae: Bombus) species. Molecular Ecology 20: 4870 ...
[94]
[PDF] IPBES Transformative Change Assessment Messages for ...
Apr 15, 2025 · • Pollinators, Pollination and Food Production (delivered 2016). • 4 Regional Assessments: the Americas, Europe and Central Asia, Africa, and ...