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Aculeata
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| Aculeata Temporal range:
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|---|---|
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| Vespula vulgaris | |
Scientific classification 
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| Kingdom: | Animalia |
| Phylum: | Arthropoda |
| Class: | Insecta |
| Order: | Hymenoptera |
| Suborder: | Apocrita |
| Infraorder: | Aculeata (but see text) |
| Superfamilies | |
Aculeata is an infraorder of Hymenoptera containing ants, bees, and stinging wasps. The name is a reference to the defining feature of the group, which is the modification of the ovipositor into a stinger. However, many members of the group cannot sting, either retaining the ovipositor, or having lost it altogether. A large part of the clade is parasitic.
This group includes all of the eusocial Hymenopterans.
The oldest aculeates are known from the Late Jurassic Karabastau Formation of Kazakhstan, represented by the family Bethylonymidae, which may be para- or polyphyletic.[1]
Classification
[edit]The use of the name Aculeata has a long history at the rank of infraorder or division. The Aculeata are a monophyletic, or good natural group, containing all the descendants of a single common ancestor. The Aculeata are therefore maintained as a taxon, either at infraorder or division rank or as an unranked clade.[2][3][4][5][6]
| Hymenoptera |
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References
[edit]- ^ Grimaldi, David A.; Peñalver, Enrique; Barrón, Eduardo; Herhold, Hollister W.; Engel, Michael S. (2019-11-07). "Direct evidence for eudicot pollen-feeding in a Cretaceous stinging wasp (Angiospermae; Hymenoptera, Aculeata) preserved in Burmese amber". Communications Biology. 2 (1): 408. doi:10.1038/s42003-019-0652-7. ISSN 2399-3642. PMC 6838090. PMID 31728419.
- ^ Johnson, Brian R.; Borowiec, Marek L.; Chiu, Joanna C.; Lee, Ernest K.; Atallah, Joel; Ward, Philip S. (2013). "Phylogenomics resolves evolutionary relationships among ants, bees, and wasps". Current Biology. 23 (20): 1–5. doi:10.1016/j.cub.2013.08.050. PMID 24094856.
- ^ Branstetter, Michael G.; Danforth, Bryan N.; Pitts, James P.; Faircloth, Brant C.; Ward, Philip S.; Buffington, Matthew L.; Gates, Michael W.; Kula, Robert R.; Brady, Seán G. (2017). "Phylogenomic Insights into the Evolution of Stinging Wasps and the Origins of Ants and Bees". Current Biology. 27 (7): 1019–1025. doi:10.1016/j.cub.2017.03.027. PMID 28376325.
- ^ Schulmeister, S. (2003). "Simultaneous analysis of basal Hymenoptera (Insecta), introducing robust-choice sensitivity analysis". Biological Journal of the Linnean Society. 79 (2): 245–275. doi:10.1046/j.1095-8312.2003.00233.x.
- ^ Schulmeister, S. "Symphyta". Archived from the original on 21 June 2010. Retrieved 28 November 2016.
- ^ Peters, Ralph S.; Krogmann, Lars; Mayer, Christoph; Donath, Alexander; Gunkel, Simon; Meusemann, Karen; Kozlov, Alexey; Podsiadlowski, Lars; Petersen, Malte (2017). "Evolutionary History of the Hymenoptera". Current Biology. 27 (7): 1013–1018. doi:10.1016/j.cub.2017.01.027. hdl:2434/801122. PMID 28343967.
External links
[edit]- Tree of Life Web Project: Aculeata
- [1] Bugguide.net: Aculeata
- Johnson, B. et al. Phylogenomics resolves evolutionary relationships among ants, bees, and wasps. NCBI
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Aculeata
View on Grokipediafrom Grokipedia
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 |
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
- https://www.antwiki.org/wiki/Nuptial_Flights_and_Mating
- https://www.antwiki.org/wiki/Myrmecochory