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Papilio polytes

Papilio polytes, commonly known as the common Mormon, is a species of in the family Papilionidae, characterized by its and female-limited of toxic butterflies such as Pachliopta aristolochiae . Native to tropical and subtropical regions of , it features a of 70–115 mm, with males displaying uniform black wings marked by white spots and females exhibiting polymorphic forms that mimic various unpalatable models for predator avoidance . The species is widespread across countries including , southern , , , the , and southwestern , inhabiting diverse environments from open forests and rainforests to agricultural fields, suburban gardens, and urban areas at elevations of 200–1,000 m . The life cycle of P. polytes spans approximately 28–30 days from egg to adult, with females laying pale yellow, globular eggs singly on host plants from the Rutaceae family, such as Citrus species (e.g., key lime and mandarin orange) and Murraya koenigii ; . Larvae, which resemble bird droppings in early instars for camouflage and later turn light green with a prominent eye-like marking, feed voraciously and deploy an osmeterium—a bifurcated, orange gland—to release defensive chemicals against predators . Pupae are either green or brown, blending with foliage, and adults emerge with males living 3–4 days and females up to 6–8 days, during which they reproduce year-round, peaking from January to April . Notable for its ecological role, P. polytes engages in puddling behavior where males aggregate at damp soil or mud to extract sodium and for reproductive benefits, while females focus on oviposition . The species' , controlled by genetic loci including the doublesex gene, allows female polymorphism with up to five distinct forms in some populations, enhancing survival through resemblance to distasteful species . Though not endangered and lacking special , P. polytes can act as a minor pest on crops due to larval feeding, yet it serves as a key model in studies of , , and ; .

Taxonomy and Nomenclature

Taxonomy

Papilio polytes is classified within the following taxonomic hierarchy: Kingdom Animalia, Phylum Arthropoda, Class Insecta, Order , Family Papilionidae, Genus Linnaeus, 1758, Species polytes Linnaeus, 1758. It belongs to the subgenus Menelaides Hübner, , which encompasses a monophyletic of approximately 70 characterized by specific wing venation and coloration patterns. The was first described by in 1758 in his , based on specimens from , and has since been placed within the genus, which comprises over 200 worldwide. Taxonomic revisions have grouped P. polytes into the polytes species group, a within Menelaides that includes five closely related exhibiting similar morphological traits and geographic distributions across . Early classifications, such as those in Talbot's 1939 The Fauna of British India, recognized extensive intraspecific variation leading to the description of numerous based primarily on regional differences in wing coloration and markings. Over 20 subspecies of P. polytes have been recognized, reflecting its wide and morphological ; notable examples include P. p. romulus Cramer, 1775, found in and characterized by distinct black wing bands, and P. p. alphenor & , 1907, occurring in with variations in hindwing spotting. Subspecies delineation traditionally relies on morphological criteria, such as differences in forewing cell markings, hindwing tail shape, and overall coloration intensity, often tied to geographic isolation across the , , and island populations. More recent genetic analyses, including mitochondrial and nuclear , have supported these divisions by revealing low but consistent divergence levels (p-distances of 1-8%) and reciprocal in some island versus mainland populations. Post-2018 molecular phylogenetic studies have confirmed the monophyly of the polytes species group using Bayesian inference on multi-locus datasets, with posterior probabilities exceeding 0.95, reinforcing P. polytes' distinct status within Papilio. A key revision in 2023 elevated Papilio javanus Oberthür, 1888 (previously a subspecies of P. polytes) to full species rank based on reproductive isolation, genetic divergence, and biogeographic separation between Sundaic island taxa and mainland Asian forms; however, as of 2025, the valid name for this species is Papilio theseus Cramer, 1777, and Papilio alphenor has also been elevated to full species status, expanding the polytes group. These updates, drawing from exhaustive sampling of 78% of Papilio species and over 50% of known subspecies, underscore the role of allopatric diversification in shaping the group's taxonomy.

Names

The scientific name Papilio polytes derives from the genus , Latin for "butterfly," and the specific polytes, from the Greek root "poly-" meaning "many," referencing the ' polymorphic female forms that mimic multiple other butterflies. This nomenclature highlights the butterfly's biological trait of female polymorphism, enabling for protection against predators. First described by in 1758 as part of the Papilionidae family within the order , the name has undergone minor revisions in taxonomic classifications. Some later authors proposed reassigning it to the genus Princeps based on morphological traits, but this is now regarded as a junior of Papilio. The common English name "Common Mormon" alludes to the female's ability to exhibit multiple morphs mimicking distasteful species, drawing an to polygamy in early Mormon culture where one male mates with many females. This name underscores the species' reproductive strategy and complex. In , it bears regional vernacular names such as "Bahurupi" in , evoking its variable appearances.

Physical Description

Males

Males of Papilio polytes typically have a of 77–96 mm, making them slightly smaller than females on average. Their dorsal wings are predominantly jet-black, adorned with a series of pale yellow or white oval spots that traverse the center of the hindwing, forming a prominent discal band; the forewings feature smaller white marginal spots along the outer edge. The hindwings include distinctive red tornal spots near the anal angle, often with a sooty-black outline, and are equipped with elongated tails characteristic of the Papilionidae family. A key sexual characteristic of males is the presence of brand-like markings on the wings, formed by specialized scales known as androconia, which disperse pheromones to attract females during . These structures are absent in females and contribute to , providing a baseline against which the ' polymorphic female forms can be compared. Subspecies variations in are subtle but notable. Overall, males remain monomorphic across the species, lacking the adaptations seen in females.

Females

Female Papilio polytes exhibit , with females generally larger than males, possessing a ranging from 87 to 105 mm compared to 77 to 96 mm in males. This increased size contributes to their broader wings and greater body mass, adaptations that support enhanced egg-laying capacity during . The baseline female form displays a darker black ground color on the wings with subtle pale yellow or white markings, including streaks between the veins on the forewings and a series of four elongated white spots on the hindwings, where the two middle spots are more prominent. Unlike males, which feature specialized wing brands or androconia for dispersal, females lack these structures, resulting in smoother wing surfaces without such scent scales. The abdomen is notably larger and more robust than that of males, providing space for developing eggs and facilitating oviposition on host plants such as species. Antennal shows minimal dimorphism, with both sexes bearing clubbed antennae typical of Papilionidae. These traits underscore the pronounced in P. polytes, where the baseline serves as the foundation for various polymorphic extensions observed in some populations.

Polymorphic Forms

The females of Papilio polytes exhibit pronounced polymorphism, manifesting in distinct morphs characterized by variations in wing coloration, patterning, and subtle differences in tail shape, which enable them to resemble locally abundant unpalatable for predator avoidance. These forms represent adaptations through , where the palatable P. polytes gains protection by imitating toxic models from the genus Pachliopta. The non-mimetic cyrus form closely resembles the male but features a paler ground color, broader white bands on the forewings, and more prominent red submarginal crescents on the hindwings, with spatulate tails similar to those of males. This form occurs throughout the ' range and is relatively common in Indian populations, such as in , where it constitutes a notable proportion of females. The stichius form, the most prevalent mimetic morph across much of the range including , mimics the common rose () with a glossy black ground color, a large irregular white patch on the forewings, white streaks on the hindwings, and vivid red spots along the hindwing margins; the tails are elongated and black-tipped. It dominates in populations from the to , often comprising over 50% of females in surveyed areas. The polytes form also mimics the common rose but with a more extensive white discal band on the forewings and reduced black scaling, resulting in brighter white areas, alongside red anal spots and similar tail structures to stichius. This variant appears in regions where the model is abundant, contributing to local frequency distributions alongside stichius. The form imitates the melanic variant of the common rose, displaying darker wings with minimized white markings, a narrow white forewing band, and subdued red hindwing spots, while retaining the characteristic swallowtail tails. It is primarily found in and Philippine populations overlapping with the model, at lower frequencies compared to stichius. The form resembles the crimson rose () in a subdued manner, with tawny-red forewings marked by black borders and veins, black hindwings featuring white submarginal spots and red basal patches, and slightly shorter, blunter tails. This morph is localized to southern and , occurring at moderate frequencies where the model is present. These polymorphic forms enhance survival by deterring predators familiar with the distasteful models, with morph frequencies varying regionally to match local model abundances and thereby optimizing anti-predator efficacy.

Gynandromorphs

Gynandromorphs of Papilio polytes are rare individuals displaying bilateral , characterized by male traits on one side of the body and traits on the other, resulting in asymmetric patterns. For instance, the right may exhibit the typical dark male coloration with pale bands, while the left shows a polymorphic form such as f. , which mimics distasteful model species like . This condition underscores the species' pronounced , where s exhibit polymorphic absent in males. The first documented case appeared in a 1930 report from a specimen collected in , described as a "freak" P. polytes with one forewing and hindwing pair resembling the male-like female form f. cyrus and the opposite side the mimetic female form f. romulus. Subsequent records include additional specimens from the , showcasing similar bilateral splits in wing coloration and pattern. These anomalies originate from chimeric development during embryogenesis, typically due to an error in sex chromosome segregation during the initial mitotic divisions, leading to tissues with differing sex determination (e.g., female cells on one side and male cells on the other). Such gynandromorphs are exceptionally rare, with fewer than 10 verified specimens recorded in and collections as of 2025, all from Asian populations.

Distribution, Habitat, and Status

Geographic Range

Papilio polytes is native to South and , with its range extending from through and eastward to , , , , , southern , , the , , (including ), and southern (particularly the ). This widespread distribution spans diverse tropical and subtropical regions, where the species is commonly observed in various landscapes. The species exhibits polytypy, with numerous adapted to regional variations across its . For instance, P. p. occurs in , , , and parts of , while P. p. polytes is found in and southern . Other notable subspecies include P. p. pasikrates in and P. p. alcindor on and nearby islands. P. p. sugama is reported in southern . These reflect local morphological and genetic adaptations within the broader Asian . Papilio polytes typically inhabits elevations from up to 1,500 m, though it has been recorded breeding at lower altitudes and as stragglers higher. A recent observation in 2024 extended its known altitudinal limit to 2,000 m in , , suggesting potential for occasional higher occurrences. As of 2025, the distribution of Papilio polytes remains stable and extensive, with no documented large-scale contractions, owing to its adaptability and common status across its range.

Habitat Preferences

_Papilio polytes, commonly known as the common Mormon, inhabits a variety of tropical and subtropical environments across its range in Asia, including open forests interspersed with meadows, tropical rainforests, and agricultural fields. This species shows a strong preference for areas where host plants such as Citrus species are available, often found at elevations between 200 and 1000 meters. It avoids strong winds exceeding 5 to 10 m/s by sheltering in undergrowth, indicating a reliance on protected, vegetated zones for survival. Within these broader habitats, P. polytes favors microhabitats that support key behaviors, such as sunny clearings for basking to regulate body temperature and proximity to sources or moist for puddling, where males congregate to extract minerals from damp , puddles, or edges. These are commonly observed in home gardens, and riparian forests, and woody plains to moist patches, where they fly low and swiftly, typically between 9 AM and 2 PM. The presence of suitable perches like shrubs and branches for resting and pupation further defines preferred microhabitats. Seasonal activity of P. polytes varies regionally, with increased abundance and breeding during wet periods; in , populations peak during the (June to September) and post-monsoon months, correlating with higher humidity and availability of fresh host plant foliage. In contrast, drier seasons see reduced immature stage densities, though the species can breed year-round in suitable conditions. This pattern aligns with observations in , where wet seasons support higher larval survival on young leaves. P. polytes demonstrates notable adaptations to human-modified landscapes, thriving in orchards, suburban gardens, and even large urban areas due to the widespread cultivation of host plants like curry leaf (Murraya koenigii) and citrus crops, which provide consistent food resources. This flexibility allows the species to persist in agricultural settings and city parks, where nectar sources such as and are also abundant, enhancing its resilience amid .

Conservation Status

_Papilio polytes, commonly known as the Common Mormon, is not listed as threatened on the and has no special , reflecting its stable and widespread populations across its range. This assessment aligns with earlier reviews, such as those in the 1985 IUCN publication on threatened swallowtail butterflies, which did not identify it as vulnerable. The species remains common and abundant, particularly in core regions like , where it is one of the most frequently observed swallowtails. Despite its overall security, P. polytes faces localized threats from habitat loss driven by and , which reduce available and breeding areas. applications on host plants, such as species, pose risks to larval stages by causing direct mortality or sublethal effects on development. Additionally, potential shifts in its range due to could impact distribution patterns, though current evidence suggests resilience given its adaptability. No targeted conservation programs are in place for P. polytes owing to its commonality, but it benefits from general protections within reserves such as India's , where habitat preservation supports its populations. In countries like , it is recognized as a common under broader initiatives. Ongoing monitoring of butterfly diversity in protected areas helps maintain its status without requiring species-specific interventions.

Mimicry and Polymorphism

Batesian Mimicry

is an antipredator strategy in which a palatable evolves to resemble an unpalatable or toxic model , thereby deceiving predators and gaining protection without the costs of . In Papilio polytes, this form of mimicry is female-limited and polymorphic, with certain female morphs imitating distasteful to evade avian predators, while males and non-mimetic females rely on other defenses such as or speed. This adaptation allows the harmless P. polytes to exploit the learned avoidance behaviors of predators toward the models, enhancing female survival in predator-rich environments. The primary models mimicked by P. polytes females belong to the genus Pachliopta, which are unpalatable swallowtails containing toxic aristolochic acids sequestered from their host plants. Specific mimetic morphs include f. polytes, which resembles Pachliopta aristolochiae (common rose); f. theseus, which mimics Pachliopta hector (crimson rose); and f. romulus, which imitates other Pachliopta species in sympatric regions. These resemblances extend to wing coloration, patterning, and even tail projections, creating a mimicry ring where multiple P. polytes morphs coexist with their models across Southeast Asia. In some populations, additional morphs like f. stichius target locally abundant Pachliopta variants, ensuring adaptive matching to regional predator pressures. Field experiments demonstrate that this mimicry effectively deters predators, with mimetic P. polytes replicas suffering approximately 30% less predation from birds compared to non-mimetic forms in controlled trials. Avian predators, such as those trained on models, exhibit reduced attack rates on mimics due to generalized avoidance of the warning signals, confirming the protective benefit. At the population level, the polymorphism stabilizes through negative , where rare morphs gain higher fitness by better matching local model abundances, maintaining mimicry ring diversity among sympatric species over evolutionary timescales.

Genetic Mechanisms

The polymorphism and female-limited Batesian mimicry in Papilio polytes are primarily regulated by the doublesex (dsx) gene through sex-specific alternative splicing, which produces distinct isoforms that direct wing pattern development exclusively in females. This mechanism ensures that male butterflies exhibit the default non-mimetic form, while females can adopt one of several mimetic morphs resembling toxic model species, as identified through genetic mapping and expression analyses. The dsx gene's female-specific isoform activates downstream regulatory networks, including genes like ebony and optix, to orchestrate pigment deposition and pattern formation on the wings. The inheritance of mimetic forms follows an autosomal dominant pattern at a single locus designated H, where dominant alleles (H) produce mimetic females and recessive homozygotes (h/h) yield non-mimetic forms, with strict sex-limitation preventing expression in males. This locus corresponds to a genomic region encompassing dsx, forming a that spans approximately 130 kb and includes structural variants such as inversions, which suppress recombination and maintain linkage between alleles controlling multiple mimetic traits. Recent / editing studies in related species, including confirmation of dsx functionality in 2025 analyses of P. alphenor (formerly a P. polytes ), have validated that targeted disruptions in this region abolish female-specific , underscoring the supergene's modular genetic architecture. Evolutionarily, the dsx in P. polytes preserves polymorphism by balancing selection on non-recombining haplotypes, allowing adaptive mimetic alleles to co-evolve without breaking apart under varying ecological pressures across populations. This non-recombining structure, characterized by high sequence divergence and selective sweeps, parallels supergenes in other species and highlights how regulation rather than multi-gene clusters drives rapid evolutionary innovation in wing patterns. Such mechanisms contribute to the species' diverse polymorphic forms by enabling stable inheritance of complex traits.

Behavior and Ecology

Adult Behavior

Adult Papilio polytes butterflies display distinct sex-specific strategies that support their reproductive needs. Males commonly participate in , aggregating at damp soil, puddles, stream edges, or even carrion and to ingest sodium and , which enhances male mating success and improves hatching rates in females receiving nuptial gifts. In contrast, females forage selectively on from a variety of flowers of short herbs and shrubs. Flight patterns in adult P. polytes are adapted for evasion and . Males and non-mimetic females employ swift, erratic flights to predators, characterized by rapid directional changes. Mimetic female morphs, however, exhibit behavioral by adopting slower, more linear flight paths with reduced directional change (mean DC ≈ 45° vs. 60° in non-mimetics) and lower variability in path deviation, closely resembling the steady flight of their unpalatable models like . This resemblance in complements their wing pattern , deterring predators more effectively. Courtship in P. polytes involves males actively seeking mates through patrolling flight or perching on branches and leaves, initiating pursuits upon detecting a . Males rely on cuticular hydrocarbons (CHCs) as contact pheromones, primarily alkenes like 7-tricosene (male-specific) and profiles dominated by C23–C25 compounds, to recognize conspecific and reject heterospecifics or males, with copulation attempts dropping significantly when CHCs are removed. During , males perform aerial chases targeting active , showing preferences that vary by context: for instance, active are courted regardless of , while stationary mimetic receive more attention in some populations. Female choice plays a minimal role, with no observed preferences based on male traits, as males are monomorphic; success hinges more on male persistence and female receptivity. Adult P. polytes lack territoriality, with males neither defending sites nor forming leks, relying instead on opportunistic . Movements are limited to short distances within home ranges, facilitated by wind currents, and the does not engage in long-distance migrations.

Life Cycle Stages

_Papilio polytes undergoes complete , consisting of four distinct life stages: , , , and . This process typically spans 28 to 38 days under tropical conditions, influenced by factors such as and . The eggs are small, spherical, and greenish-yellow, measuring 0.94 to 1.3 mm in diameter, and are laid singly on the upper surfaces of tender host plant leaves. They turn pale yellow to brownish-black as they mature, with an incubation period of 3 to 4 days at temperatures around 29°C. The larval stage comprises five s, lasting a total of 14 to 25.5 days. Early instars (first to fourth) exhibit bird-dropping , appearing as pale brown or dark with white saddle markings and spines for against predators. In the fifth instar, the shifts to a smooth dark green form with white and black sinuous bands and prominent eyespots, growing up to 50 mm in length. Defense involves eversion of the , a bifurcated organ that emits a foul odor when threatened. Pupation occurs after a brief prepupal period, resulting in a chrysalis that measures 25 to 32 mm and is suspended by a silk girdle. The exhibits dimorphism, appearing green or brown to blend with foliage or stems, and features typical papilionid head projections. The pupal stage lasts 10 to 11 days, after which the adult emerges. The overall cycle duration shortens with higher temperatures in tropical environments.

Host Plants and Interactions

Larval Host Plants

The larvae of Papilio polytes primarily utilize plants from the family as host plants, with a strong preference for species such as Citrus limon (lemon) and Citrus maxima (pomelo), which provide essential foliage for development. Other notable hosts include , a wild shrub that supports larval growth in natural settings. These plants contain secondary metabolites like , which the larvae detoxify enzymatically to avoid during feeding. Larvae deploy an —a bifurcated, —to release endogenous defensive chemicals against predators. Feeding behavior is instar-specific, with early instars targeting tender young leaves to minimize mechanical resistance and optimize nutrient intake, while later instars consume more mature foliage. This selective consumption supports efficient development and contributes to the bird-dropping that camouflages the early instars visually. Host range variations occur across regions, incorporating species (prickly-ash) as alternative wild hosts in certain areas. Such flexibility aids to local flora but can extend to cultivated areas. Ecologically, P. polytes larvae pose a significant threat to agriculture, as gregarious feeding leads to severe defoliation of young shoots and entire trees during outbreaks, impacting production in orchards. This status underscores the species' role in balancing herbivore-plant dynamics while challenging commercial cultivation.

Adult Nectar Sources

Adult Papilio polytes rely on floral as their primary energy source, providing carbohydrates and crucial for flight, , and production. This exhibits opportunistic feeding , visiting a broad array of flowering without strong specialization, which enhances its adaptability across diverse habitats from forests to urban gardens. Observations indicate that adults prefer flowers with accessible , often those featuring tubular corollas or dense clusters that allow efficient probing with their . Studies in tropical regions have documented P. polytes nectaring on up to 104 plant species, predominantly (38 species) and shrubs (43 species), with a preference for cultivated perennials that bloom in dense masses. Key families include (21 species), (9 species), and (11 species), reflecting the butterfly's attraction to nectar-rich inflorescences. For instance, (Asteraceae) and Mikania cordata (Asteraceae) are frequently visited due to their abundant, small flowers, supporting high nectaring activity with over 500 recorded events in garden settings. In agricultural ecosystems, P. polytes has been observed foraging on 51 plant species across 24 families, with , , and being prominent. Representative examples include Lantana camara (), an invasive shrub offering continuous nectar throughout the year; Tithonia diversifolia (), valued for its bright, composite flowers; and Asystasia gangetica (), a common weed providing reliable blooms from July to December. Other favored sources encompass Ixora spp. () and Jatropha curcas (), which feature elongated floral structures suited to the butterfly's feeding apparatus. These interactions not only sustain the adults but also aid in pollinating weedy and cultivated plants in human-modified landscapes.

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    [PDF] 55 FORAGING BEHAVIOUR OF SOME PAPILIONID BUTTERFLIES ...
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    [PDF] 40 Study on nectar plants of few butterfly species at agriculture ...
    Nectar is a food for many adult butterflies visit plants with specific coloured ... Papilio polytes; Ga: Graphium agamemnon,. Plant type: We: Weed, Cp ...