In entomology, an alate refers to a winged reproductive form of social insects, such as ants, termites, and certain aphids, which emerges from mature colonies to facilitate dispersal and reproduction.[1] These individuals, often larger than other colony members, possess fully developed wings and compound eyes, enabling flight during swarming events.[2] The term also functions as an adjective to describe any insect or structure bearing wings or wing-like extensions, contrasting with apterous (wingless) forms.[3]Alates play a critical role in the life cycle of eusocial insects by promoting genetic diversity and colony expansion. In ants and termites, they are produced seasonally—typically in spring or summer—when colonies reach maturity, and they depart en masse in nuptial flights to mate with alates from unrelated colonies, reducing the risk of inbreeding.[4] Following mating, the female alate sheds her wings; in ants, the male typically dies soon after, while in termites, both shed their wings and the mated pair (queen and king) excavates a chamber in soil or wood to lay eggs, initiating a new colony.[5] This reproductive strategy allows species like the eastern subterranean termite (Reticulitermes flavipes) to colonize distant areas, though most alates perish without successfully founding a colony.[6]Beyond social Hymenoptera and Isoptera, alates appear in other insects like aphids, where winged forms develop in response to environmental cues such as overcrowding, enabling long-distance migration and the transmission of plant pathogens.[3] In botany, "alate" describes seeds, leaves, or stems with wing-like flanges that aid in wind dispersal, as seen in maple seeds.[7] The presence of alates, particularly indoors, often signals a nearby mature infestation requiring investigation.[8]
Etymology and General Definition
Etymology
The term "alate" derives from the Latin adjective alātus, meaning "winged," which is formed from āla, denoting "wing" or "winglike structure."[9][10] This Latin root traces back to the Proto-Indo-European aks-, signifying "axis" or "joint," particularly in reference to the pivotal joint of a wing or arm.[9]The word entered English in the 1660s through scientific literature, with the earliest recorded use in 1661 by naturalist Robert Lovell in his descriptions of natural history.[11] Initial biological applications appeared in mid-17th-century entomological texts to describe insects possessing wings, and by 1668, the term had extended to botanical contexts for plant structures with winglike extensions, such as winged stems or petioles.[11][12]An archaic adverbial sense of "alate" meaning "lately" or "of late" dates to the late 15th century and stems from Middle English "a- + late", unrelated to and dismissed in modern scientific parlance.[10] The term's adoption in formal biological nomenclature solidified in the 18th century with the rise of binomial systems, where it became a standard descriptor in species epithets to indicate winged or winglike morphology, as seen in names like Passiflora alata.[11]
General Definition
In biology, "alate" primarily serves as an adjective describing structures or organisms that possess wings or winglike extensions, such as the functional wings of insects or the flattened, wing-shaped appendages on plant seeds that aid in dispersal.[7] As a noun, it refers to a winged individual within a species exhibiting polymorphism, particularly the reproductive forms that develop wings for mating and colony founding in social insects.[13] This term is distinct from its occasional non-biological uses, focusing instead on analogous or true wing structures in the animal and plant kingdoms.[14]The word's scope encompasses both zoological applications, where it denotes fully developed, functional wings enabling flight, and botanical contexts, where it applies to winglike modifications like alate petioles or samaras.[15] In modern taxonomy, "alate" is commonly incorporated into species epithets to indicate winged characteristics, as seen in Dioscorea alata (greater yam), where "alata" highlights winglike features on tubers or stems.[16] It contrasts with the antonym "apterous," meaning wingless, which describes forms lacking such structures.[17]Pronounced /ˈeɪ.leɪt/ in English, "alate" derives briefly from the Latin ala meaning "wing," and archaic variants like "alated" are sometimes encountered in older texts, though "winged" remains a common synonym in contemporary usage.[11][18]
In Entomology
Definition and Characteristics
In entomology, the term alate (from Latin alatus, meaning "winged") refers to a winged adult insect, particularly the reproductive forms in social species such as ants, termites, and aphids. These alates are typically the sexually mature individuals that emerge from colonies to engage in dispersal and mating, contrasting with wingless (apterous or apterous) castes like workers or soldiers.[1]Key characteristics of alates include fully developed wings—forewings and hindwings of equal length in termites and unequal in ants—allowing powered flight during nuptial swarms. They often possess larger compound eyes for navigation, ocelli (simple eyes) in some species, and a more robust, pigmented body compared to non-reproductive forms, which aids in desiccation resistance and visual signaling. Alates are generally larger than workers; for example, termite alates measure 1/4 to 1/2 inch (6-13 mm) in length including wings, with soft-bodied, pale to dark coloration depending on the species. In ants, alate queens and males exhibit elbowed antennae, while termite alates have straight, beaded antennae, aiding in species identification during swarms. These traits evolve to support long-distance dispersal, with wings composed of chitinous membranes supported by veins for structural integrity during flight.[19][20][5]Unlike the passive wing-like structures in botany, entomological alates enable active locomotion via muscle-powered wingbeats, typically at frequencies of 10-20 Hz in termites and higher in ants, facilitating escape from mature colonies and colonization of new habitats. This polymorphism—winged vs. wingless forms—arises from environmental or genetic cues, allowing colonies to balance reproduction and maintenance. Taxonomically, alate morphology is crucial for identifying reproductive castes in keys and descriptions, such as distinguishing alate ants from similar wasps by waist segmentation.[2]
Role in Insect Reproduction and Dispersal
In eusocial insects such as ants and termites, alates function as the primary dispersers of gametes, enabling sexual reproduction and colony propagation. Alate queens (gynes) and males emerge from mature colonies to participate in nuptial flights, where they mate in mid-air or at aggregation sites, often with partners from unrelated colonies. This mating process allows fertilized queens to found new colonies independently, carrying stored sperm for lifelong egg production while males typically die post-copulation.[21][22][23]The winged morphology of alates facilitates long-distance dispersal, with flight capabilities varying by species but often spanning tens to hundreds of meters on average, and up to several kilometers in favorable conditions such as wind assistance. For instance, in fire ants (Solenopsis invicta), alates can travel up to 12-16 km over water, promoting gene flow across populations and reducing inbreeding depression by mixing genetic material from distant colonies. This dispersal strategy contrasts with worker-mediated budding in some species, emphasizing alates' role in independent colony establishment over local expansion.[24][25][23]Alate production is typically triggered by environmental cues like seasonal changes in photoperiod, temperature, and humidity, or internal colony signals such as overcrowding and resource depletion, signaling optimal conditions for dispersal. In ants, mature colonies allocate resources to produce alates seasonally, often synchronizing flights across populations to maximize mating success. However, this investment incurs high energetic costs, as developing alates require substantial nutrition—up to 60% of a queen's body mass in reserves—and represent only a small proportion of colony output (e.g., 1-5% in many ant species), reflecting the trade-off between growth and reproduction. Mortality during production and flight exceeds 99% due to predation and physiological demands, ensuring alates are produced judiciously when dispersal benefits outweigh risks.[21][23][26]Ecologically, alate dispersal enhances biodiversity by enabling colonization of unoccupied habitats and maintaining metapopulation dynamics in fragmented landscapes. By introducing genetic variation, alates support adaptive evolution in response to environmental pressures, while their flights integrate colonies into broader food webs as prey for birds, bats, and other predators. However, this vulnerability to predation during dispersal phases regulates population densities, preventing overexploitation of resources and influencing communitystructure in insect societies.[23][21]
Examples of Alate Insects
In the order Hymenoptera, alate forms are prominent in ants, where queens and males develop wings for nuptial flights. For instance, in the red imported fire antSolenopsis invicta, alate queens and males swarm from mature colonies during warm spring or summer evenings to mate, after which the females shed their wings and establish new colonies, while wingless workers remain the primary foragers and defenders within established nests.[27]Termites in the order Isoptera (now classified under Blattodea) also exhibit alate reproductives that play a key role in colony founding. In species such as Reticulitermes flavipes and R. hageni, alates emerge en masse during swarming events, typically in spring or fall on warm, humid days following rain, to disperse and pair for mating; these winged forms differ from ant alates in their straight antennae and equal-length wings, and they shed wings by chewing them off at the base rather than simply dropping them.[5][28]Among Hemiptera, aphids demonstrate remarkable polymorphism, producing alate morphs adapted for long-distance migration. In the black bean aphid Aphis fabae, alate females are generated parthenogenetically under conditions of crowding, poor host quality, or seasonal cues, enabling host alternation between winter hosts like spindle trees and summer crops such as beans, thus facilitating range expansion and escape from predators.[29][30]Thrips in the order Thysanoptera show similar dispersal polymorphism. The western flower thripsFrankliniella occidentalis produces fully winged (macropterous) adults that actively fly or are wind-assisted for dispersal to new hosts, contrasting with short-winged (brachypterous) forms that remain on crowded plants; this variation allows the pest to invade greenhouses and field crops globally.[31][32]Alate polymorphism extends to other Hymenoptera, such as bees, where male drones in species like the honey beeApis mellifera develop wings specifically for orientation and mating flights to drone congregation areas during the reproductive season. In Psocoptera, known as booklice or barklice, many species exhibit wing dimorphism, with alate individuals dispersing from deteriorating habitats to colonize new substrates, while apterous forms dominate stable environments, highlighting adaptive flexibility across both eusocial and solitary insects.[33][34]
Dealation
Dealation refers to the process by which alate insects, particularly reproductive females in social species like ants, shed their wings after mating, transitioning to a wingless (dealate) state suited for sedentary colony founding.[35] This shedding typically involves autotomy, where the queen uses her mandibles or legs to break the wings at their basal articulation points, a mechanism facilitated by a weakened attachment that allows easy detachment without significant injury. In some cases, enzymatic processes contribute to the degradation of wing remnants post-shedding, though the primary mechanism is mechanical removal.[36]The process occurs shortly after the nuptial flight and mating, often triggered by hormonal shifts including elevated juvenile hormone levels that promote reproductive maturation and inhibit further flight.[37] In ant queens, such as those of Solenopsis invicta, mating introduces neurotransmitters like tyramides from males, which disinhibit dealation and initiate physiological changes including wing loss.[38] Males typically die soon after mating due to exhaustion and lack of further reproductive roles, while virgin queens in colonies are prevented from dealating by inhibitory pheromones from the mated queen until dispersal.[36] The shed wings are often consumed by the queen or discarded, marking the end of the dispersive phase.Physiological adaptations in dealate queens include the histolysis (breakdown) of indirect flight muscles, which are resorbed to provide proteins and energy for oogenesis and early colony establishment, exemplifying a trade-off between dispersal and reproduction. In species like Lasius niger and Atta cephalotes, this muscle resorption can supply up to 30% of the energy needed for the first brood, allowing the queen to survive without external food during claustral founding.[39] The now-redundant wings and associated structures are no longer maintained, redirecting resources to nest excavation and egg production within the colony.Dealation ensures the queen's commitment to a sedentary lifestyle, focusing colony efforts on internal reproduction and foraging by workers rather than further dispersal. This behavior enhances colonystability in social insects, as the wingless queen cannot easily abandon the nest. Fossil evidence from Cretaceousamber (approximately 99 million years ago) includes dealate queens of stem-group ants like Gerontoformica, indicating that dealation and associated social adaptations were already present in early eusocial Hymenoptera, supporting the rapid evolution of complex colony structures.[35]
In Botany
Definition and Characteristics
In botany, the term alate serves as an adjective to describe flattened, wing-like expansions on various plant parts, such as ribs, keels, fruits, seeds, stems, or petioles, which resemble wings in form but possess no motility or flight capability. These structures are integral to plant morphology, where they denote adaptations that primarily aid in dispersal, especially via wind (anemochory), by increasing surface area to facilitate passive transport.[40][41][42]Key characteristics of alate structures include their composition from corky or papery tissues, often derived from the cork cambium or thin epidermal extensions, providing lightweight yet resilient forms; for example, the prominent wings on stems of Euonymus alatus form as corky ridges that enhance visual and structural distinction. These expansions are frequently lignified, incorporating secondary cell walls with lignin for added durability against mechanical wear and environmental exposure during dispersal. In terms of variation, alate features typically measure 1-10 cm in length or width, scaling with the host organ, and are distinctly absent in wingless (non-alate) counterparts, such as unwinged fruits in species exhibiting polymorphic dispersal strategies.[43][44][45]Unlike the active, chitinous wings of insects used for powered flight in entomology, botanical alates are immobile appendages designed solely for passive aerodynamic assistance in seed or fruit dissemination, highlighting an evolutionary convergence in wing-like morphology to exploit wind currents for anemochory without requiring metabolic energy for locomotion.[46][47]Taxonomically, alate is employed in formal descriptions to specify morphological traits, such as "fructus alatus" for fruits bearing wing-like appendages, which aids in distinguishing species and genera; this usage contrasts alate conditions in seeds (e.g., samaras) from those on stems or petioles, enabling precise classification in floras and keys.[40][44]
Types of Alate Structures
Alate structures in plants manifest in diverse forms depending on their location and tissue composition, primarily serving morphological adaptations. These features often arise from modifications in the outer layers of plant tissues, such as the epidermis or periderm.[42]Alate fruits and seeds commonly include samaras, which are single-seeded dry fruits characterized by lateral wings formed by thin, membranous extensions of the pericarp surrounding the achene.[48] Another variant consists of achenes enclosed within persistent, wing-like calyces that extend outward from the floral structure, creating a winged appearance around the seed.[49]Alate stems and branches typically feature corky flanges or ridges that develop along the length of the stem, often through longitudinal splitting of the bark or activity of cork cells in the periderm.[42] Decurrent wings represent a specific subtype, where tissue from adjacent leaf bases extends downward along the stem, forming continuous ridge-like projections.[50]Alate petioles and leaves exhibit expanded bases or margins, where the petiole broadens into wing-like expansions that connect to the leafblade or stem, potentially enhancing surface area in certain environments.[42] In arid-adapted species, these expansions can contribute to water storage within the petiole tissues.[51]Composite types of alate structures include hybrid forms such as alate capsules observed in families like Bignoniaceae, where the fruit wall develops flattened or ridged extensions alongside winged seeds.[52] These composite features often originate from modifications in the periderm for stem-related wings or the epidermis and underlying tissues for fruit and seed structures.[42]
Examples in Plants
In the genus Acer (family Sapindaceae), alate samaras are a prominent feature, consisting of paired wings that encircle the seed body, as seen in species like the Norway maple (Acer platanoides). These structures, typically 3–5 cm long, arise from paired carpels and mature in pendulous clusters during late spring to summer in temperate regions of Europe and North America.[53][54]The winged spindle (Euonymus alatus, family Celastraceae) exemplifies alate stems, bearing four prominent corky wings along young branches that expand to 5–10 mm wide, providing a distinctive ridged appearance. Native to temperate forests in East Asia, including China, Japan, and Korea, this deciduousshrub reaches 4–6 m in height and is widely cultivated ornamentally for its vibrant red autumn foliage and compact form.[55][56]Ash trees in the genus Fraxinus (family Oleaceae) produce alate samaras with a single terminal wing attached to one side of the seed, differing from the paired configuration in maples; for instance, the European ash (Fraxinus excelsior) features samaras 3–4 cm long that mature in autumn. This temperate species, distributed across Europe and western Asia, supports wide-ranging seed distribution through these lightweight, indehiscent fruits.[57][58]Beyond temperate examples, the tipu tree (Tipuana tipu, family Fabaceae) from subtropical South America showcases alate pods resembling samaras, with a single broad wing spanning 6–7 cm around the flattened legume. This tropical canopy tree, reaching 20–30 m, produces these structures in abundance following its bright yellow inflorescences, contributing to its ornamental use in warmer climates.[59][60]In arid environments, desert shrubs of the genus Atriplex (family Amaranthaceae), such as fourwing saltbush (Atriplex canescens), display alate bracts that form four papery wings around the utricle, measuring 1–2 cm across and aiding adaptation to saline soils. These halophytic species thrive in salt deserts of North America and beyond, where the winged bracts enclose the seed while enhancing tolerance to high salinity levels up to 25 dS/m.[61][62]Alate structures thus illustrate botanical diversity, appearing in temperate lineages like Acer, Euonymus, and Fraxinus across Eurasia and North America, as well as in tropical and arid taxa such as Tipuana in South America and Atriplex in global drylands, reflecting adaptations suited to varied climates.[63]
Ecological Significance
Alate structures in plants primarily facilitate anemochory, the wind-mediated dispersal of diaspores, by augmenting surface area and promoting autorotation or gliding, which prolongs airborne duration and enables seeds to travel distances typically ranging from 10 to 100 meters or more, thereby minimizing intraspecific competition near the parent plant.[64][65] This adaptation is particularly advantageous in open habitats where wind currents can carry samaras, such as those of Acer species, far from the source, enhancing colonization of new areas.[66]Beyond dispersal, alate features serve protective roles, including deterrence of herbivores through structural modifications like spiny or toughened wings on fruits and stems that hinder feeding or oviposition by insects.[67] In xerophytic environments, winged stems contribute to water conservation by altering architecture to reduce exposure to desiccating winds and support stability against gusts, while also potentially limiting transpiration through modified surface morphology.[68] These functions collectively bolster plant survival in arid or exposed ecosystems by integrating mechanical defense with physiological efficiency.Evolutionarily, alate structures exemplify convergent evolution across disparate lineages, such as the fin-winged fruits observed independently in families like Asteraceae and Fabaceae, driven by selective pressures for enhanced dispersal in fragmented or windy landscapes.[44]Fossil evidence from the Eocene epoch reveals early angiosperms with alate fruits, including samara-like forms in genera akin to modern Ailanthus, indicating that these adaptations arose during a period of rapid diversification amid changing paleoclimates around 50 million years ago.[69]In contemporary ecosystems, alate traits face conservation challenges from climate change, as projected reductions in wind speed and altered patterns could curtail dispersal efficacy for anemochorous species, impeding migration to suitable habitats.[70] Conversely, these structures facilitate the spread of invasive species, exemplified by the winged samaras of maples like Acer platanoides, which enable prolific wind dispersal and rapid establishment in non-native regions, exacerbating biodiversity threats.[71]