Fact-checked by Grok 2 weeks ago

Aculeata

Aculeata is a monophyletic clade within the order Hymenoptera, comprising the ants, bees, and stinging wasps, and is defined by the evolutionary modification of the female ovipositor into a sting apparatus used for venom injection and defense.^[1] This diverse group encompasses over 70,000 described species across approximately 31 families, making it one of the most species-rich lineages in the Hymenoptera.^[2] Taxonomically, Aculeata includes several superfamilies, notably Chrysidoidea (including tiphiid and scoliid wasps), Vespoidea (encompassing vespid and mutillid wasps), Formicoidea (ants), and Apoidea (comprising apoid wasps and bees).^[2] Recent phylogenomic analyses, based on ultraconserved elements, have resolved key relationships within the clade, confirming that ants (Formicidae) are the sister group to the Apoidea, with bees nesting deeply within the paraphyletic sphecoid wasps.^[2] The clade originated in the Late Jurassic to Early Cretaceous around 150–140 million years ago, with major diversifications linked to the radiation of flowering plants and the evolution of eusociality in multiple lineages.^[3] Aculeates exhibit a wide range of life histories, from solitary predation and parasitism to advanced eusocial colonies, and play pivotal ecological roles as pollinators (especially bees), predators of pest insects (many wasps and ants), and soil engineers (ants).^[2] Their sting, derived from the ovipositor, serves primarily for subduing prey or defense, though it is absent or vestigial in males and workers of some social species.^[1] The group's ecological and economic importance is immense, with bees alone responsible for pollinating a significant portion of global agriculture, while ants and wasps contribute to natural pest control and biodiversity maintenance.^[2]

Taxonomy

Definition and Characteristics

Aculeata is a monophyletic infraorder within the suborder Apocrita of the order Hymenoptera, encompassing ants, bees, and stinging wasps, and distinguished from the sawfly suborder Symphyta by the presence of a constricted petiole and a modified ovipositor adapted for stinging rather than egg-laying or plant tissue penetration.^[4]^[2] The defining synapomorphy of Aculeata is the transformation of the ancestral ovipositor into an aculeus, a venom-injecting stinger that serves as a potent defensive and predatory tool, marking a key evolutionary innovation within Hymenoptera.^[5]^[6] Key characteristics of Aculeata include the "wasp waist," formed by a narrow petiole connecting the mesosoma and metasoma, which provides flexibility for locomotion and stinging maneuvers, alongside a modified female reproductive tract featuring venom glands and reservoirs integrated with the stinger apparatus.^[4]^[7] The stinger is present only in females, though functionality varies across groups, with secondary reductions in some lineages such as certain ants and bees.^[5]^[7] This group represents a diverse radiation of over 100,000 described species across approximately 40 families, with estimates suggesting total diversity could exceed 300,000 undescribed species.^[8]^[9]

Classification

Aculeata comprises an infraorder of the order Hymenoptera, within the suborder Apocrita, encompassing ants, bees, and various stinging wasps characterized by a modified ovipositor into a venomous sting apparatus. The group is divided into 8 superfamilies, reflecting a hierarchical structure informed by both morphological and molecular data. Key superfamilies include Chrysidoidea (encompassing families such as Bethylidae and Chrysididae, with jewel wasps as notable examples), Pompiloidea (including Pompilidae, the spider wasps), Tiphioidea, Thynnoidea, Vespoidea (featuring Vespidae, such as yellowjackets and hornets, along with Mutillidae), Formicoidea (dominated by the family Formicidae, the ants), Apoidea (including apoid wasps like those in Crabronidae and the bees in Anthophila, such as Apidae and Megachilidae), plus minor groups Sierolomorphoidea and Scolioidea.^[10]

Superfamily	Key Families and Examples	Approximate Described Species
Chrysidoidea	Bethylidae (bethylid wasps), Chrysididae (cuckoo wasps, jewel wasps)	~5,000
Pompiloidea	Pompilidae (spider wasps)	~5,000
Tiphioidea	Tiphiidae (tiphiid wasps)	~1,000
Thynnoidea	Thynnidae (thynnid wasps)	~2,000
Vespoidea	Vespidae (yellowjackets, hornets), Mutillidae (velvet ants)	~12,000
Formicoidea	Formicidae (ants)	~16,000
Apoidea	Crabronidae (digger wasps), Apidae (bees, including honeybees), Megachilidae (leafcutter bees)	~30,000 (including ~21,000 bees)
Other (Sierolomorphoidea, Scolioidea)	Various minor families	~3,000

Aculeata includes over 120,000 described species worldwide as of 2024, with estimates suggesting total diversity could exceed 300,000 when accounting for undescribed taxa; Apoidea accounts for approximately 30,000 species (of which bees comprise about 21,000), Formicidae around 16,000, and Vespoidea roughly 12,000.^[8]^[11]^[12] Recent taxonomic revisions have incorporated molecular phylogenomic data, such as ultraconserved element (UCE) analyses, leading to reclassifications including the confirmation of bees as a derived lineage within Apoidea and the elevation of certain groups like Sapygidae to family status within Pompiloidea. As of 2024, the classification recognizes 8 superfamilies based on integrated phylogenomic studies. These updates resolve previous uncertainties in superfamily boundaries and family-level relationships.^[2]^[13]

Phylogeny

Aculeata occupies a key phylogenetic position as the sister group to Ichneumonoidea within the suborder Apocrita of the order Hymenoptera, a relationship corroborated by comprehensive phylogenomic analyses that integrate molecular data from hundreds of taxa across the order. This placement highlights the evolutionary transition from parasitoid lifestyles in Ichneumonoidea to the predatory and stinging behaviors characteristic of aculeates. Seminal work by Branstetter et al. (2017) utilized ultraconserved elements (UCEs) from 854 loci, yielding over 203,000 base pairs of aligned sequence data from 187 aculeate species, to resolve higher-level relationships with high statistical support across maximum-likelihood, Bayesian, and coalescent-based methods. Within Aculeata, the phylogeny delineates a basal grade comprising superfamilies such as Chrysidoidea (e.g., cuckoo wasps) and Pompiloidea (e.g., spider wasps), which represent early-diverging lineages characterized by solitary hunting strategies and lack of eusociality. The core Aculeata forms a well-supported monophyletic clade that includes Formicidae (ants) as the sister group to a combined Apoidea (bees and apoid wasps) plus Vespoidea (vespoid wasps), a topology consistently recovered in the Branstetter et al. study with bootstrap values exceeding 95% and posterior probabilities near 1.0. Apoidea itself is monophyletic, with bees (Anthophila) emerging as a derived, monophyletic subgroup nested within the paraphyletic assemblage of sphecoid wasps (e.g., Crabronidae and Sphecidae), underscoring the evolutionary origin of bees from within a diverse radiation of ground-nesting wasps. This nested structure of bees within sphecoid wasps has been further reinforced by targeted phylogenomic efforts focusing on Apoidea. Key phylogenomic studies, including those employing transcriptomic data from hundreds of genes, provide evidence for the divergence of major aculeate clades during the Late Jurassic to Early Cretaceous. For instance, Peters et al. (2017) analyzed transcriptomes from 185 hymenopteran species to estimate divergence times, placing the last common ancestor of ants and Apoidea in the Late Jurassic (mean 162 Ma, 95% CI 192–136 Ma). Recent studies as of 2024, such as those integrating fossil-calibrated clocks, confirm crown-group Aculeata origins around 160–190 million years ago. These findings resolve longstanding controversies, such as the debated origins of ants—previously hypothesized to be closely related to hunting vespoid wasps—and the precise placement of bees relative to other wasps, which were uncertain due to reliance on morphological data alone. Recent DNA sequencing approaches, particularly those integrating large-scale nuclear gene datasets, have definitively settled these debates by demonstrating ants' basal position within the core Aculeata and bees' derivation from sphecoid ancestors, with no significant conflicts across analytical methods.^[2]^[14]

Evolution

Fossil Record

The fossil record of Aculeata begins in the Late Jurassic, with the oldest known specimens recovered from the Karabastau Formation in southern Kazakhstan, dated to approximately 165 million years ago (Ma). These compression fossils include members of the extinct family Bethylonymidae, considered basal aculeates, which exhibit primitive morphological features such as reduced wing venation and ovipositor structures indicative of early stinging wasp lineages.^[15] Mesozoic diversity expanded notably during the Cretaceous, particularly in amber deposits from Myanmar (Burmese amber), dated to around 99 Ma. These inclusions preserve primitive bees such as Melittosphex burmensis (family Melittosphecidae), marking one of the earliest known anthophilous (flower-visiting) aculeates with branched hairs and pollen-collecting adaptations. Early ants, including Sphecomyrma freyi from New Jersey amber (~100 Ma), reveal the onset of formicoid morphology and potential eusocial behaviors through preserved worker castes.^[16] Compression fossils from Early Cretaceous sites, such as the Lulworth Formation in England (~145–140 Ma), further document Pompilopterus species, aiding classification via detailed wing venation patterns that distinguish aculeate from non-stinging hymenopterans.^[15] The Cenozoic era shows a marked expansion of aculeate lineages, with Eocene amber from the Baltic region (~45 Ma) yielding diverse modern families including Vespidae (e.g., eumenine and vespine wasps) and Apidae (eusocial bees like meliponines). These amber fossils preserve fine details of social structures, such as caste dimorphism in ants (Formicidae) and pollen-laden corbiculae in bees, indicating established eusociality.^[17] By the Oligocene (~34–23 Ma), fossils from sites like Enspel in Germany include eusocial bees such as Apis henshawi, with open-nesting behaviors inferred from body size and wing morphology, alongside ant colonies showing advanced division of labor.^[18] Preservation in compression fossils from sedimentary layers, particularly in Jurassic and Cretaceous deposits, has been crucial for aculeate systematics, as wing venation—featuring reduced crossveins and aculeate-specific sclerites—allows precise taxonomic placement despite incomplete bodies.^[15] Amber inclusions from Myanmar and the Baltic provide exceptional three-dimensional preservation, revealing soft tissues and behaviors that compression fossils often obscure.

Evolutionary Origins and Diversification

Aculeata, the clade encompassing stinging wasps, bees, and ants, originated from parasitoid ancestors within the Apocrita suborder of Hymenoptera during the Late Jurassic, approximately 165–150 million years ago (Ma).^[19] These early aculeates evolved from wasps that used their ovipositors to lay eggs in or on host insects, a strategy that shifted toward prey immobilization as the ovipositor modified into a venom-delivering stinger, enabling active hunting rather than solely parasitism.^[3] This transition marked a pivotal adaptation, allowing aculeates to subdue larger prey and defend against threats, setting the stage for their ecological expansion.^[5] Key innovations further propelled aculeate evolution, including the development of eusociality in ants, bees, and certain wasps, which facilitated cooperative brood care, division of labor, and colony-based foraging from solitary hunting lifestyles.^[20] Eusociality, characterized by overlapping generations and reproductive altruism, likely arose multiple times within Aculeata, enhancing survival through collective defense and resource efficiency.^[21] Concurrently, bees underwent co-evolution with angiosperms, transitioning to pollination mutualisms that provided reliable nectar and pollen resources, driving specialized floral adaptations and vice versa.^[22] Diversification occurred in distinct phases: an initial basal radiation during the Late Jurassic established core lineages, followed by a Cretaceous explosion linked to the angiosperm radiation beginning ~125 Ma, which opened new niches for pollinators and predators.^[15] Cenozoic peaks, particularly in tropical regions during the Eocene and Miocene, amplified species richness through habitat fragmentation and climatic shifts.^[23] Major drivers included ecological opportunities such as insect predation and pollination interactions, which buffered against environmental stresses.^[24] The Cretaceous-Paleogene (K-Pg) mass extinction ~66 Ma had minimal impact on aculeates due to their small body size, diverse diets, and subterranean or colonial habits, allowing rapid post-extinction recovery and further radiation.^[25]

Morphology and Anatomy

Sting Apparatus

The sting apparatus in Aculeata represents a key synapomorphy, derived from the modified ovipositor of female Hymenoptera, enabling envenomation of prey or for defense.^[26] It consists of a cuticular complex comprising two paired stylets, known as lancets (the 1st valvulae), and an unpaired central shaft, or stylet (the 2nd valvula), which together form the piercing structure.^[27] These components are supported by sclerites including the 1st and 2nd valvifers, and are integrated with associated musculature for movement.^[28] The apparatus also includes venom glands and a reservoir, where venom is produced by epithelial cells in the glands and stored in a sac connected to the sting via a duct.^[27] Venom composition across Aculeata typically includes bioactive peptides, enzymes such as phospholipases, and biogenic amines like histamine, serotonin, acetylcholine, and dopamine, which induce paralysis, pain, or tissue damage in envenomated targets.^[29] For instance, in social wasps, peptides such as vespid mastoparans contribute to membrane disruption and pain signaling.^[30] The venom reservoir, often a thin-walled sac, holds small volumes (e.g., approximately 1-2 mg in honeybee workers), sufficient for multiple injections.^[27] Modifications to the sting vary significantly among aculeate groups. In bees of the family Apidae, such as the honeybee Apis mellifera, the lancets bear multiple backward-facing barbs (about 10 per lancet), which facilitate tissue penetration but cause the sting to lodge irreversibly, leading to autotomy and the death of the worker bee as the apparatus continues pumping venom post-detachment.^[27] This barbed structure contrasts with smoother variants in some solitary bees, though linkage to pollen-collecting adaptations like the corbicula occurs in social species. In ants (Formicidae), the sting is fully developed in queens and workers of many subfamilies but reduced or absent in males, which lack the ovipositor-derived structure entirely; ant stings often feature lancets with serrated distal ends for efficient venom delivery into prey.^[31] In wasps of the family Vespidae, the sting shaft exhibits barbs on the lancets (typically 2-12 per side), with greater barb density and strength in some social species, enhancing retention during defensive stings but increasing autotomy risk.^[32] These barbs differ from the smoother, unbarbed stings in many solitary wasps, optimizing for repeated use. Functionality relies on coordinated musculature for insertion and venom injection. The lancets reciprocate via protractor (e.g., M198) and retractor (e.g., M199) muscles attached to accessory plates, driving the stylets up to 1.3 mm into tissue while a venom duct flap regulates dosage through muscular pumping.^[27] This mechanism allows precise envenomation, with barbs reducing insertion force and amplifying extraction resistance. Variations include loss or reduction in certain parasitic groups; for example, in velvet ants (Mutillidae), females are apterous and possess a prominent sting—the longest relative to body size among aculeates—while males are winged and stingless, reflecting sexual dimorphism.^[33] Conversely, in spider wasps (Pompilidae), the sting is elaborated for paralyzing large spider prey, featuring specialized flaps in the venom duct for rapid, accurate dosing and robust musculature enabling quick strikes without barbs for reusability.^[34]

Body Structure and Adaptations

Aculeates exhibit a distinctive body plan typical of Apocrita within Hymenoptera, divided into three primary tagmata: the head, mesosoma, and metasoma. The head is a compact capsule bearing large compound eyes laterally, three ocelli dorsally in most species, and paired antennae that arise from membranous sockets; in ants (Formicidae), the antennae are characteristically elbowed or geniculate at the scape-flagellum joint, facilitating sensory functions such as chemoreception. The mesosoma, comprising the fused thorax and propodeum, supports three pairs of legs and, in winged forms, the membranous wings, which display reduced venation in more derived groups like bees and ants compared to basal wasps. The metasoma, representing the posterior abdomen, is connected to the mesosoma by a constricted petiole—often forming a narrow "wasp waist"—which enhances flexibility and maneuverability.^[4] Internally, aculeates possess specialized structures adapted for their lifestyles, including strong, sclerotized mandibles used for nest excavation, prey manipulation, or resource processing across taxa. In bees (Apoidea), the foregut includes a crop, or honey stomach, serving as a storage reservoir for nectar during foraging, expandable to hold up to 50 microliters and preventing digestive enzymes from mixing with collected liquids. Leg modifications are prominent: bees often feature corbiculae or pollen baskets on the hind legs for transporting pollen, while many wasps (Vespoidea) have tarsal spines or rake-like structures on the forelegs for grooming or prey handling. These adaptations reflect the group's diversity in feeding and nesting strategies.^[4]^[35] Key adaptations in aculeates include a wide size range, from approximately 1 mm in minute ants like those of the genus Carebara to 50 mm in large spider wasps such as tarantula hawks (Pompilidae: Pepsis), allowing exploitation of varied ecological niches. Coloration often involves aposematic patterns, such as black-and-yellow stripes in many wasps, which serve as Müllerian mimicry signals to deter predators by advertising chemical defenses. Flight is powered by asynchronous indirect flight muscles in the mesosoma, enabling high wingbeat frequencies—up to 230 Hz in some bees—for efficient hovering and rapid escape, with wing coupling via hamuli ensuring coordinated motion. The metasoma tip integrates the sting apparatus in females, but the overall body form prioritizes agility and sensory acuity.^[4]^[36]^[37] Sexual dimorphism is pronounced in aculeates, with females generally larger and equipped with a functional sting derived from the ovipositor, enhancing defense and oviposition capabilities, whereas males are typically smaller, lack a sting, and possess fully developed wings for dispersal and mating. This dimorphism supports eusocial structures in many species, where female workers forgo reproduction but retain morphological traits for colony maintenance.^[4]

Biology and Behavior

Reproduction and Life Cycle

Aculeates exhibit haplodiploid sex determination, in which males develop from unfertilized haploid eggs and are thus haploid, while females develop from fertilized diploid eggs and are diploid.^[38] This system is characteristic of the entire suborder Hymenoptera, including all Aculeata.^[38] Some species, such as the Cape honeybee (Apis mellifera capensis), also display thelytokous parthenogenesis, where unmated females produce diploid female offspring through automixis, allowing worker reproduction in queenless colonies.^[39] The life cycle of aculeates involves complete metamorphosis with four distinct stages: egg, larva, pupa, and adult. Eggs are typically laid by females in nests or on provisioned prey, hatching into legless, grub-like larvae that are fed by adults—either progressively in social species or via pre-provisioned paralyzed prey in solitary ones.^[4] Larvae undergo several molts before pupating within a cocoon or sealed cell, where metamorphosis occurs, eventually emerging as winged or wingless adults depending on caste and sex. Mating behaviors vary across aculeates but often involve pheromones for attraction. In social species like bees and ants, nuptial flights synchronize the release of virgin queens and males, facilitating aerial mating swarms.^[40] Solitary wasps, such as those in the Sphecidae, typically mate near nests, with females provisioning burrows or cells by paralyzing prey using their sting before laying eggs on or near it.^[41] Reproductive strategies differ markedly between solitary and social forms. In eusocial aculeates like ants and bees, reproduction is caste-based, with queens laying most eggs, sterile female workers performing non-reproductive tasks, and haploid males (drones in bees) focused on mating.^[38] Some sphecoid wasps exhibit subsocial care, where females guard nests and provide progressive provisioning to larvae over extended periods, representing an intermediate step toward greater social complexity. Haplodiploidy promotes eusociality in these groups through kin selection, as sisters share 75% of their genes on average, favoring altruistic worker behavior.^[42]

Sociality and Foraging

Aculeata exhibit a wide spectrum of social organization, ranging from solitary lifestyles to highly advanced eusocial colonies. Most wasps, such as digger wasps in the family Sphecidae, are solitary, with females provisioning nests individually by hunting prey to feed their larvae.^[20] In contrast, primitively eusocial species occur in some bees, like those in the family Halictidae, where small colonies feature flexible roles and occasional queen succession.^[20] Advanced eusociality, characterized by distinct castes including queens, workers, and sometimes soldiers, is prevalent in ants (Formicidae), honeybees (Apidae), and yellowjackets (Vespidae), enabling large, perennial colonies with division of labor.^[43] This diversity in sociality is influenced by factors such as haplodiploid sex determination, which promotes altruism through asymmetric relatedness among siblings.^[44] Foraging strategies in Aculeata vary by group and reflect their ecological niches. Wasps and ants predominantly engage in carnivorous foraging, actively hunting insects or scavenging protein sources to provision nests or feed colonies.^[45] Bees, conversely, focus on herbivory and nectarivory, collecting pollen and nectar from flowers to support brood development and colony energy needs.^[46] Nest construction complements these behaviors, with solitary wasps using mud or soil to build individual burrows, while social species like paper wasps (Vespidae) fashion nests from chewed wood fibers and ants excavate underground galleries.^[45] Communication enhances foraging efficiency; for instance, honeybees employ the waggle dance to convey the direction, distance, and quality of food sources to nestmates.^[47] Ants and some wasps utilize pheromone trails to recruit foragers to rich resources.^[48] Defense mechanisms are integral to sociality and foraging, particularly in eusocial species. Alarm pheromones trigger rapid colony responses, such as mass stinging in honeybees or aggressive recruitment in ants and wasps, deterring predators during foraging expeditions.^[49] Territoriality is prominent in ants, where foragers mark and defend foraging trails against intruders using chemical cues and physical confrontations.^[50] In social wasps, venom-released pheromones coordinate attacks on threats near nests or food sites.^[51] Variations in these behaviors include kleptoparasitism, observed in cuckoo bees (e.g., Nomadinae) and cuckoo wasps (Chrysididae), where females invade host nests to lay eggs on provisioned resources, sparing the effort of foraging or nest-building themselves.^[52] This parasitic strategy exploits the foraging investments of solitary or eusocial hosts without contributing to colony maintenance.^[53]

Diversity and Distribution

Species Richness

Aculeata encompasses over 70,000 described species worldwide, representing a substantial portion of hymenopteran biodiversity, with estimates suggesting the true total, including undescribed taxa, could exceed 300,000 given the order's overall projected diversity of up to 1 million species. This richness is concentrated in tropical regions, where environmental complexity supports high speciation rates across ants, bees, and wasps.^[54] Among the major families, Formicidae (ants) includes approximately 16,000 described species, dominating in terms of biomass and ecological impact despite comprising about 23% of Aculeata diversity.^[11] Apidae, the largest bee family, accounts for around 5,700 species, encompassing social groups like honey bees and bumble bees that play key roles in pollination. Crabronidae stands out as the most species-rich among wasp families, with over 9,000 described species, many of which are solitary ground-nesters adapted to diverse habitats. Patterns of endemism are pronounced in isolated regions, such as Australia, where bulldog ants (Myrmecia spp.) represent a primitive lineage with over 90 endemic species exhibiting unique predatory behaviors. In Madagascar, high endemism is evident among bees, including the subspecies Apis mellifera unicolor, a tropical honey bee adapted to island conditions and contributing to local pollination networks.^[55] Habitat loss poses significant threats, with studies indicating that 10-20% of assessed wild bee species in regions like Europe face extinction risk primarily from deforestation and land conversion, a pattern likely amplified globally for Aculeata.^[56] Diversity hotspots highlight regional variations, with the Neotropics harboring exceptional wasp richness, including thousands of species in families like Vespidae and Pompilidae due to the area's floral and prey abundance. The Paleotropics, encompassing Southeast Asia and sub-Saharan Africa, serve as a center for ant diversity, where Formicidae species outnumber other Aculeata groups and drive ecosystem processes like seed dispersal.^[57]

Global Distribution and Habitats

Aculeata exhibit a cosmopolitan distribution, occurring across all major biogeographic realms on Earth except Antarctica, where bees and other groups are absent due to the lack of suitable terrestrial habitats. Ants (Formicidae) are particularly ubiquitous on landmasses, present in virtually all non-polar terrestrial ecosystems from boreal forests to arid zones. Bees (Apoidea) and wasps (Vespoidea and others) similarly span from high Arctic latitudes—such as bumblebees (Bombus spp.) foraging in tundra regions—to equatorial tropics, with wasps showing notable diversity in desert environments, including nocturnal species like those in the subfamily Sphaeropthalminae. This broad range reflects their adaptability to extreme climates, though overall diversity peaks in tropical regions. Habitat preferences among aculeates vary by taxon and lifestyle, with eusocial species like many ants and some bees forming colonies in forested areas, often nesting in rotting wood or soil rich in organic matter. Solitary nesters, including numerous wasps and ground-nesting bees, favor open grasslands and meadows for burrow construction and prey/forage access. Urban and disturbed habitats have seen successful colonization by adaptable species, such as certain ants and bees that exploit human-modified landscapes for nesting and resources. Along altitudinal gradients, aculeate diversity often increases with elevation in temperate zones, as observed in the Kyushu Central Mountains of Japan, where species richness of bees and wasps rises from lowlands to montane forests due to varied microhabitats and reduced competition. Biogeographic patterns reveal ancient origins influencing modern ranges, with some ant lineages tracing Gondwanan ancestry, as seen in ponerine ants that originated and diversified across southern supercontinent fragments before spreading globally. Bees display prominent Holarctic radiations, exemplified by carpenter bees (Xylocopa) that underwent northward expansions from Eocene origins, adapting to temperate and boreal environments. Island endemism is pronounced in isolated systems, such as the Hawaiian yellow-faced bees (Hylaeus spp.), with over 60 species unique to the archipelago and reliant on native vegetation for nesting and foraging. Dispersal in aculeates is generally limited to short-range flight or colony budding, restricting natural long-distance migration for most species. However, human-mediated introductions have dramatically expanded ranges, notably for the Argentine ant (Linepithema humile), which has achieved worldwide distribution through global trade, invading Mediterranean, subtropical, and temperate regions and altering local ecosystems.

Ecological and Human Significance

Roles in Ecosystems

Aculeates play pivotal roles in ecosystems as pollinators, predators, and decomposers, contributing to food web stability and nutrient cycling. Bees, primarily within the Apiformes, serve as the dominant pollinators for approximately 80% of the world's flowering plants, facilitating reproduction through pollen transfer while foraging for nectar and pollen.^[58] This service supports biodiversity among angiosperms and underpins about 35% of global food production by aiding pollination of 87 major crop species.^[59] Wasps, including various Aculeata families, act as secondary pollinators by incidentally transferring pollen on their bodies during nectar feeding, enhancing plant reproduction in diverse habitats despite their less specialized morphology compared to bees.^[60] In predation and biological control, many aculeate wasps and ants function as parasitoids and predators, regulating populations of herbivorous insects and other arthropods. Social wasps prey on pest species such as lepidopteran larvae, reducing their abundance in natural and agricultural settings and thereby preventing outbreaks that could disrupt plant communities.^[41] Ants similarly control pest insects through direct predation and interference, while species like spider wasps (Pompilidae) target and paralyze spiders, naturally limiting arachnid populations and maintaining balance in arthropod food webs.^[61] Additionally, ants act as soil engineers by excavating nests that aerate soil, improve water infiltration, and enhance nutrient availability, which promotes plant growth and ecosystem productivity.^[62] Aculeates also contribute to decomposition and symbiotic processes that recycle nutrients and disperse seeds. Certain wasps scavenge carrion and organic matter, accelerating breakdown and returning essential nutrients to the soil, which supports microbial activity and soil fertility.^[63] Ants facilitate myrmecochory, a form of seed dispersal where they transport seeds with lipid-rich elaiosomes to their nests, consuming the appendage and discarding the viable seed in nutrient-enriched waste piles, thereby aiding plant colonization and forest regeneration across ecosystems.^[64] Across trophic levels, aculeates occupy diverse niches: bees primarily as herbivores consuming floral resources, wasps and many ants as carnivores preying on other insects, and omnivorous behaviors emerging in social colonies where members exploit multiple food sources.^[65] These varied roles position aculeates as keystone taxa in many habitats, where their absence could cascade through food webs, reducing biodiversity and altering community structure.^[41]

Interactions with Humans

Aculeates provide substantial agricultural benefits through honey production, pollination, and biological control. The western honey bee, Apis mellifera, supports global apiculture, with managed colonies numbering approximately 102 million worldwide as of 2023 and producing about 1.9 million metric tons of honey annually as of 2023.^[66] This industry generates significant income for rural communities and contributes to products like beeswax used in cosmetics and food processing.^[67] Additionally, aculeate wasps, such as the paper wasp Polistes satan, serve as effective predators of crop pests like lepidopteran larvae, aiding biological control in agriculture without relying on chemical pesticides.^[68] Pollination by aculeates, particularly bees, underpins a vast economic value, enhancing crop yields for fruits, vegetables, and nuts, with global services estimated at $235–577 billion annually.^[69] However, interactions also pose health risks; stings from bees, wasps, and ants can trigger anaphylaxis in allergic individuals, leading to 50–100 deaths per year in the United States alone.^[70] Invasive aculeates like the red imported fire ant, Solenopsis invicta, exacerbate threats by damaging crops, livestock, and equipment, causing approximately $8 billion in annual economic losses across infested regions.^[71] Culturally and economically, aculeates inspire innovations such as swarm robotics, modeled after ant collective behaviors for tasks like resource foraging and obstacle navigation, as seen in NASA's Ant-Inspired Swarmie robots.^[72] Yet, pesticides like neonicotinoids contribute to declines, including colony collapse disorder in honey bees, where exposed colonies exhibit up to fourfold increases in parasite levels and reduced reproduction.^[73] Conservation efforts address these issues, with about 25% of native bee species imperiled in North America due to habitat loss and chemicals; responses include habitat restoration programs and bans on neonicotinoids in the European Union since 2018, alongside U.S. restrictions on high-risk uses. Recent surveys indicate record-high losses, with 55.6% of managed colonies lost in the US between April 2024 and April 2025.^[74]^[75]^[76]^[77]

References

[1]
[PDF] Bees, ants, and stinging wasps (Aculeata) - TimeTree.org
Aculeate wasps are characterized by the modification of the ovipositor into a sting. All other Hymenoptera deposit their eggs through their ovipositor, while.
[2]
Phylogenomic Insights into the Evolution of Stinging Wasps and the ...
Apr 3, 2017 · Phylogenomic Insights into the Evolution of Stinging Wasps and the Origins of Ants and Bees ... Download: Download Acrobat PDF file (1MB).
[3]
[PDF] Hymenoptera of the world: An identification guide to families.
... wasp waist"); this connection is actually the greatly narrowed articulation of abdominal terga. 1 and 2. • A true, primitive ovipositor with three pairs of.
[4]
(PDF) Functional morphology and evolution of the sting sheaths in ...
Oct 18, 2021 · The sting of the Aculeata or stinging wasps is a modified ovipositor; its function (killing or paralyzing prey, defense against predators) ...
[5]
Functional morphology and evolution of the sting sheaths in ...
Sep 17, 2019 · The sting of the Aculeata or stinging wasps is a modified ovipositor- its function (killing or paralyzing prey, defense against predators) ...Missing: definition transformation
[6]
Aculeata - an overview | ScienceDirect Topics
Aculeata refers to the species Pereskia aculeata, a member of the Cactaceae family, known for its height of up to 10 meters and its use in traditional ...
[7]
Functional anatomy of the worker honeybee stinger (Apis mellifera)
Jun 24, 2023 · The honeybee stinger is a powerful defense mechanism that combines painful venom, a subcutaneous delivery system, and the ability to autotomize.Missing: monophyly transformation
[8]
The radiation of Hymenoptera illuminated by Bayesian inferences ...
May 5, 2025 · Stinger wasps (Aculeata) are estimated to have originated during the Jurassic (e.g., Bethylonymoidea: median 171.91 Ma, CI: 164.01–183.99 Ma), ...
[9]
The abundance, biomass, and distribution of ants on Earth | PNAS
Ants are highly diverse, comprising more than 15,700 named species and subspecies (26) and possibly as many undescribed ones (27).
[10]
Evolution and systematics of the Aculeata and kin (Hymenoptera), with emphasis on the ants (Formicoidea: †@@@idae fam. nov., Formicidae) | bioRxiv
Summary of each segment:
[11]
Phylogeny of Asilidae Inferred from Morphological Characters of ...
The oldest fossils that can be unambig- uously assigned to the ... sion fossils from the Karabastau Formation of southern Kazakhstan (Late Jurassic,.
[12]
The First Mesozoic Ants - Science
Two worker ants preserved in amber of Upper Cretaceous age have been found in New Jersey. They are the first undisputed remains of social insects of Mesozoic ...
[13]
Monophyly and extensive extinction of advanced eusocial bees
Comprehensive study of bees fossilized in Baltic amber has revealed an unsuspected middle Eocene (ca. 45 million years ago) diversity of eusocial bee lineages.Missing: Vespidae | Show results with:Vespidae
[14]
Fossil insects from the Late Oligocene Enspel Lagerstätte and their ...
Nov 19, 2009 · It can be inferred that the Oligocene A. henshawi was eusocial and probably had an open-air nest like A. dorsata and other basal species of Apis ...
[15]
Evolutionary History of the Hymenoptera - ScienceDirect.com
Apr 3, 2017 · Chrysidoids are confirmed as the sister group of all remaining Aculeata [19]. We corroborate the artificial nature of the former superfamily “ ...
[16]
Key innovations and the diversification of Hymenoptera - Nature
Mar 3, 2023 · Aculeata was estimated with an age of 142 Ma (node 26; Fig. 2), placing the origin of the modified ovipositor-stinger at the edge of the ...Missing: transformation | Show results with:transformation<|control11|><|separator|>
[17]
Life History and the Transitions to Eusociality in the Hymenoptera
Obligate eusociality meets the definition of a major evolutionary transition, and such transitions have occurred five times in the Hymenoptera.
[18]
THE EVOLUTION OF EUSOCIALITY - PMC - NIH
It is evident that bees, and also wasps, are springloaded, that is, strongly predisposed with a trigger, for a rapid shift to eusociality, once natural ...
[19]
Direct evidence for eudicot pollen-feeding in a Cretaceous stinging ...
Nov 7, 2019 · By the time of the main period of angiosperm radiation, some 120–90 Ma, aculeate wasps were well evolved.
[20]
The origins of species richness in the Hymenoptera: insights from a ...
Apr 27, 2010 · There are four insect orders with over 100,000 described species, of which the Hymenoptera is one [23–25]. Little work has addressed the ...
[21]
The Angiosperm Terrestrial Revolution buffered ants against extinction
Mar 11, 2024 · Ants maintained their diversification trajectory throughout the Oligocene and Miocene (≈1.2 and ≈1.45 times the background net ...
[22]
The angiosperm radiation played a dual role in the diversification of ...
Jan 22, 2024 · We found that angiosperms played a dual role that changed through time, mitigating insect extinction in the Cretaceous and promoting insect origination in the ...
[23]
(PDF) Fossil Social Insects - ResearchGate
Sep 28, 2023 · The evolution of eusociality is one of the major transitions in the history of life, particularly in the insects. Now, fossil termites and ants ...
[24]
Morphology of sting apparatus of Chrysidoidea (Hymenoptera ...
Oct 26, 2020 · The main synapomorphy for Aculeata is the sting apparatus, which allows the female to envenom potential prey or hosts.Missing: defense subdual
[25]
Morphology of sting apparatus of Chrysidoidea (Hymenoptera ...
The main synapomorphy for Aculeata is the sting apparatus, which allows the female to envenom potential prey or hosts. The sting is the modified ovipositor ...
[26]
Functional and Proteomic Insights into Aculeata Venoms - PMC
Mar 16, 2023 · Aculeate venoms are mixtures of peptides, enzymes, biogenic amines, and other organic compounds, such as formic acid [32,33,34,35,36,37].
[27]
A review of the venom peptides of social wasps (Hymenoptera ...
... amines are acetylcholine, histamine, serotonin, dopamine, noradrenaline, and adrenaline. These amines play different roles in the biological effects of the ...
[28]
(PDF) Structure of the sting apparatus and associated exocrine ...
In the sting the lancets have at the distal end a serrated surface, which may be important to inject venom into the prey. The venom gland has secretory cells ...
[29]
Comparative Morphology of the Stinger in Social Wasps ...
Aug 14, 2021 · Both solitary and social wasps have a fully functional venom apparatus and can deliver painful stings, which they do in self-defense.
[30]
Stinging wasps (Hymenoptera: Aculeata), which species have ... - NIH
May 2, 2018 · We find that velvet ants (Hymenoptera: Mutillidae) have the longest sting compared to their body size out of any bee, wasp, or ant species.Missing: waist petiole glands
[31]
Functional morphology of the sting apparatus of the spider wasp ...
An unpaired flap in the venom duct provides a dose of venom in the sting. This mechanism probably enhances the speed and accuracy of the wasp's sting movements.
[32]
Honey Bee Anatomy - Ask A Biologist
Jun 13, 2017 · A storage sac, used in honey bees to carry nectar. The honey stomach is hardened to prevent fluids from entering the body at this location. ...
[33]
Proportions of aposematic colouration in bees and wasps ...
Feb 28, 2025 · This group comprises over 70,000 described species, divided into approximately 40 families ( Michener 2007 ; Broad and van Noort 2024 ).
[34]
Paradoxes of Hymenoptera flight muscles, extreme machines - NIH
Feb 23, 2022 · Here we review unusual properties of flight muscles (FM) of bees and wasps. We discuss how adaptations to support extreme rates of FM contraction have evolved.
[35]
Single locus complementary sex determination in Hymenoptera
Jan 5, 2006 · All members have haplodiploid sex determination; males are haploid (one chromosome set) and females are diploid (two chromosome sets).
[36]
Inheritance of thelytoky in the honey bee Apis mellifera capensis
Jan 14, 2015 · Curiously, asexual reproduction via thelytokous parthenogenesis (the production of diploid females without fertilisation) is widespread in one ...
[37]
Nuptial Flights and Mating - AntWiki
Jun 9, 2023 · Ant mating includes female-calling where females call males, and male-aggregation where males gather at specific sites for females to fly into.Missing: Aculeata provisioning
[38]
Ecosystem services provided by aculeate wasps - Wiley Online Library
Apr 29, 2021 · We identified 962 plant species, of diverse taxonomy, which depend either solely or partially on wasps for their pollination (Fig. 4). These ...I. Introduction · Ii. Regulating Services · (1) Regulation Of Natural...
[39]
Does haplodiploidy help drive the evolution of insect eusociality?
Mar 15, 2023 · In his seminal paper, Hamilton (1964) proposed that the evolution of sociality was especially frequent within the order Hymenoptera because the ...
[40]
Evolution of the neuronal substrate for kin recognition in social ...
Aug 1, 2023 · A comprehensive view on the evolution of social recognition in Hymenoptera and its implication in the repeated emergence of eusociality is crucial.
[41]
Haplodiploidy and the Evolution of Eusociality: Worker Revolution
Hamilton suggested that inflated relatedness between sisters promotes the evolution of eusociality in haplodiploid populations. Trivers and Hare observed ...
[42]
SOCIAL WASP (HYMENOPTERA: VESPIDAE) FORAGING BEHAVIOR
Reflecting these learning abilities, the behavior of foraging wasps is influenced by cues that are associated with prey but do not emanate directly from the ...
[43]
[PDF] Convergent Evolution of Food Recruitment Mechanisms in Bees and ...
In this chapter I compare and briefly, though not exhaustively, summarize what is known about food recruitment in bees and social wasps. The focus is on recent ...
[44]
The Functional Significance of the Honeybee Waggle Dance - jstor
But, as soon as the hive is rotated back to a vertical position, both the wag- gle dances and outward flight paths again become correctly oriented. c) In the " ...
[45]
[PDF] A REVIEW ON: THE ANT FORAGING AND COMMUNICATION
ABSTRACT: Ants show a wide range of foraging strategies and this activity is tightly regulated irrespective of the mode of recruitment used.
[46]
Alarm Pheromones—Chemical Signaling in Response to Danger
Aug 7, 2025 · Two common alarm postures in Formicidae. Formicid ants respond to alarm pheromone with either aggressive or escape behaviors.
[47]
Foraging by predatory ants: A review - PubMed
Oct 21, 2024 · In this review, we show that predatory ants have a wide range of foraging behavior, something expected given their phylogenetic distance and ...Missing: Aculeata bees wasps<|control11|><|separator|>
[48]
Pheromones in Social Wasps - ScienceDirect.com
In this chapter, we reviewed the main chemical substances which mediate recognition between colony members and coordinate nest defense, alarm and recruitment.
[49]
Low Host Specialization in the Cuckoo Wasp, Parnopes grandior ...
Feb 20, 2020 · Within the Aculeata, brood parasitoids (females laying eggs on or into the host immatures), kleptoparasites (females laying eggs on the host ...
[50]
[PDF] Origins, evolution, and diversification of cleptoparasitic lineages in ...
Other lineages of cleptoparasitic Aculeata exhibit strategies con- sistent with those seen in bees. Members of the wasp family Pom- pilidae prey on spiders. Two ...
[51]
Evolution and systematics of the Aculeata and kin (Hymenoptera ...
Although crucial to the general evolutionary conception of the Aculeata, the character state "ovipositor modified as sting" was deleted from this matrix as ...
[52]
Formicidae (ants, fourmis) | INFORMATION - Animal Diversity Web
Approximately, 12,000 species of ants ( Formicidae ) are documented worldwide; however, scientists estimate that closer to 15,000 actually exist. Within this ...
[53]
Apidae - Wikipedia
Apidae is the largest family within the superfamily Apoidea, containing at least 5700 species of bees. The family includes some of the most commonly seen ...Nomadinae · Apinae · Euglossini · Xylocopinae
[54]
Crabronidae - Wikipedia
The Spheciformes included well over 200 genera, containing well over 9000 species. Revision of these taxa resulted in the restriction of the Sphecidae ...
[55]
Myrmecia (ant) - Wikipedia
These ants are commonly known as bull ants, bulldog ants or jack jumper ants, and are also associated with many other common names. They are characterized by ...Taxonomy and evolution · Behaviour and ecology · Interaction with humans
[56]
(PDF) Genetic diversity of the endemic honeybee: Apis mellifera ...
Aug 5, 2025 · Apis mellifera unicolor is a tropical honeybee endemic of Madagascar. Comprehensive knowledge about its mitochondrial and nuclear genetic ...
[57]
Mounting risks threaten survival of wild European pollinators - IUCN
Oct 11, 2025 · An estimated 10% of wild bees in Europe (at least 172 out of 1,928 assessed species) are at risk of extinction. That compares to 77 threatened ...
[58]
(PDF) Systematics and diversity of Neotropical ants - ResearchGate
Apr 16, 2021 · The Neotropical region has 137 genera and around 3,100 species of ants. The Neotropics seem to have been the “cradle” and “museum” of the groups ...Missing: hotspots Paleotropics
[59]
New global map of ant biodiversity reveals areas that may hide ...
Aug 3, 2022 · New global map of ant biodiversity reveals areas that may hide undiscovered species | University of Michigan News.Missing: Aculeata Neotropics wasps Paleotropics
[60]
Bee Pollination - USDA Forest Service
Pollinators are responsible for assisting over 80% of the world's flowering plants to reproduce including animals that assist plants with their reproduction ...
[61]
Pollinators and Global Food Security: the Need for Holistic Global ...
May 9, 2016 · Globally, 87 of major food crops depend on animal pollination. Together these account for 35 % of the world food production volume. Pollinator ...On Pollinators And Their... · Global Pollinator Decline... · Loss Of Habitat, Foraging...
[62]
Debunking wasp pollination: Wasps are comparable to bees in ...
Apr 9, 2024 · Wasps are comparable to bees in plant interactions, body pollen, and single-visit pollen deposition, showing they are effective pollinators.Missing: secondary | Show results with:secondary
[63]
Spider Wasp | NC State Extension - Entomology
Value in Pest Management Spider wasps contribute to natural control of spider populations. However, because they parasitize spiders, that themselves are ...
[64]
Ants and termites increase crop yield in a dry climate - PMC - NIH
Mar 29, 2011 · Various studies have shown that ants and termites help to create soil structure, influence aeration, water infiltration and nutrient cycling in ...
[65]
Wasps are a gardener's friend | UMN Extension
These wasps help the world's ecosystems function, as wasps pollinate flowers, control pests, spread seeds and help decompose carcasses. ... scavenge for ...
[66]
Myrmecochory - AntWiki
Jun 2, 2024 · Seed dispersal by ants is typically accomplished when foraging workers carry diaspores back to the nest, after which the elaiosome is removed or ...Terminology · 2019 · 2018
[67]
The conservation value of Aculeata communities in sand quarries ...
This concerned mostly herbivores (19.29% of the guild) and parasitic species (23.81% of the guild). For the cumulated Aculeata community, the probability of ...
[68]
World Bee Day 2025: Africa honey production has highest global ...
Globally, honey production reached 1 894 thousand tonnes in 2023, with Africa having the highest growth rate.
[69]
Global Managed Honeybee Population Exceeds 100 Million Colonies
The world's managed honeybee population has reached approximately 102.1 million colonies as of 2023, containing an estimated 1.4-4.8 trillion individual ...
[70]
The Economic Importance of Bee$, BeeSpotter, University of Illinois
The value of pollination by bees is estimated around $16 billion in the US alone. We would be unable to enjoy most of our favorite fruits, vegetables, or nuts ...Introduction · Economic Influence · Economic Problems · Problem Trends
[71]
Social wasps are effective biocontrol agents of key lepidopteran ...
Nov 6, 2019 · We provide evidence that the social paper wasp Polistes satan is a successful predator on the larvae of two economically important and resilient crop pests.
[72]
Economic value of Pollinators | Bayer Global
Jan 10, 2025 · Pollination from honey bees, native bees, and flies deliver billions of dollars (US) in economic value. Between $235 and $577 billion (US) worth of annual ...
[73]
Health Risks of Bee and Wasp Stings - Division of Research Safety
Mar 18, 2022 · According to IDPH, about one percent of the U.S. population is allergic to bee and wasp venom, and about 50 to 100 people die each year from ...
[74]
Red Imported Fire Ant | National Invasive Species Information Center
Imported fire ants have invaded about 350 million acres in the U.S., and they cause approximately $8 billion in economic damages per year. Learn more about ...
[75]
Nasa's Ant-Inspired Swarmie Robots
Sep 14, 2016 · The behavior of the robot swarm mimics the central-place foraging strategy of ants to find and collect resources in a previously unmapped ...Missing: biomimicry | Show results with:biomimicry
[76]
Colony Collapse Disorder: An Incomplete Puzzle
There was up to a fourfold increase in the levels of Nosema in honey bees from the imidacloprid-exposed colonies, regardless of whether 5 or 20 ppb were fed. “ ...
[77]
[PDF] Pollinators in Peril - Center for Biological Diversity
More than half (749) of native bee species with sufficient data are declining, and nearly 1 in 4 (347) are imperiled, with 24% in serious peril.
[78]
Bees and pesticide regulation: Lessons from the neonicotinoid ...
As a result, the use of neonicotinoids in open-field crops is now banned in the EU and restricted in some areas of the US, Philippines and Canada.
[79]
EPA Actions to Protect Pollinators
Temporarily halted the approval of new outdoor neonicotinoid pesticide uses until new bee data are submitted and pollinator risk assessments are complete.