Papilio polytes

Geographic Range

Common mormon butterflies (Papilio polytes) are widely distributed throughout Asia. They are found in Pakistan and southern Asia, including India and all regions east of it, Malaysia and Indonesia, and on the coast of southern China, the Philippine islands, and the southwestern islands of Japan. (Sehgal, et al., 2009; Smetacek, 2006; Suwarno, et al., 2010a; Suwarno, et al., 2010b; Suwarno, 2010)

Habitat

Common mormon butterflies are adaptable, inhabiting tropical and subtropical areas, open forests interspersed with meadows, tropical rainforests, agricultural fields, suburban gardens, and even large cities. Common mormon butterflies cannot fly in winds that blow more quickly than 5 to 10 m/s; thus, they cannot inhabit windy areas that lack sheltering undergrowth. Common mormon butterflies need access to water or moist earth. The males often engage in puddling behavior, wherein they drink salty fluids from moist ground, puddles, stream edges, coastlines, carrion, or feces. Common mormons require access to the plants that comprise their adult diet and the diet of their larvae. Also necessary is the availability of branches or twigs for pupal support. Common mormons have been found at elevations of 200 to 1000 m, though they may not be limited to that altitudinal range. (Arun, 2008; Barua, et al., 2004; Rangana, 2010; Sehgal, et al., 2009; Suwarno, et al., 2010b; Suwarno, 2010; Van Lien and Yuan, 2003; Watanabe, 1979)

Physical Description

Papilio polytes eggs are globular and a pale yellow. As they age, the eggs become light brown. The appearance of common mormon eggs is similar to the eggs of common lime butterflies, to the point that distinguishing the eggs of these two species is difficult. However, common lime butterflies lay their eggs on the axial part of the leaf, the stems, or on the thorns, while common mormons do not.

As a first-instar caterpillar, P. polytes has a distinctive head characterized by conspicuous sutures (shallow grooves), primary setae (large, spiky projections), and hypognathous mandibles (the lower mandible is longer than the upper mandible). Its body is light brown with white splotches, causing the caterpillar to resemble bird droppings (which serves as good camouflage). On the prothorax is an osmeterium, which is a forked, fleshy organ that emerges from the anterior of the caterpillar when it is threatened and secretes noxious chemicals to ward off predators. The caterpillar grows bigger but looks similar as it progresses through the fourth instar. In its fifth instar, the larva becomes light green, and a broad, raised stripe appears near its head.

The pupa of P. polytes is either light green, with two yellow-green patches on either side of the pupal case, or light brown.

Adult common mormon butterflies have a wingspan of 70 to 115 mm. Their hindwings have swallowtails. Adult males are smaller than females. All common mormon males look the same, with black wings that are dotted with a row of white spots across the hindwings and on the margins of the forewings. The upper forewing also has a row of white spots, which get smaller toward the wing tip. Common mormons occasionally have red crescents on the margins of each wing.

Adult females have several polymorphisms, most of which mimic other butterfly species that are unpalatable. The cyrus form is non-mimetic. It is very similar in appearance to males but has paler coloration and more distinct red crescents. The cyrus form is found across the common mormon's entire range. The most common is the stichius form, which mimics Pachliopta aristolochiae (common rose butterfly) and is found in common mormon populations from the Himalayas to Japan, Sulawesi, and Sri Lanka. The theseus form mimics the common rose butterfly's black form and is found where the habitat of both butterflies overlaps in parts of Indonesia and the Philippines. The romulus form mimics Pachliopta hector (crimson rose butterfly) but is duller in color. These mimetic polymorphisms, which effectively disguise the common mormon butterfly as other, toxic butterfly species, enable common mormon females to avoid being eaten by predators. Because P. polytes is palatable and mimics unpalatable butterfly species, this predator evasion strategy is an example of Batesian mimicry. (Durgesh, et al., 2011; Kunte, 2013; Rangana, 2010; Sehgal, et al., 2009; Smetacek, 2006; Suwarno, et al., 2010a; Tan, 2011; Vasait, 2002)

  • Sexual Dimorphism
  • female larger
  • sexes colored or patterned differently
  • female more colorful
  • Range wingspan
    80 to 115 mm
    3.15 to 4.53 in

Development

Common mormon butterflies lay about 20 to 25 eggs on the undersides and upper surfaces of leaves. Their preferred host plants typically belong to the family Rutaceae and include cultivated citrus plants such as the key lime tree and the mandarin orange. The oviposition preference of females on host plants is important, because the nutrients obtained by Papilio polytes when it is a larva greatly affect the length of larval development, the length of the adult butterfly's life, and the number of eggs that an adult female can lay.

The development of P. polytes from egg to adult takes 28 to 30 days. Both in the wild and in captivity, the eggs typically hatch after 3 to 5 days. The larva eats its egg shell before commencing to feed on the leaves of the host plant. Papilio polytes has five larval instars, or periods between molts. Larvae develop for about 14 to 20 days before pupating.

The pupa hangs by an attachment to a branch or twig, and a silk harness further supports the pupal case. Papilio polytes pupates for about 10 days, after which its emergence takes 15 to 18 minutes. After 40 to 50 minutes, the butterfly crawls up a dry twig and pumps its wings to circulate blood through its capillaries and dry its wings. After 60 to 75 minutes, the adult butterfly flies away. (Durgesh, et al., 2011; Hori, 2003; Rangana, 2010; Sehgal, et al., 2009; Shobana, et al., 2010; Smetacek, 2006; Suwarno, et al., 2010a; Tan, 2011; Vasait, 2002)

Reproduction

Papilio polytes males both patrol for females and use a waiting technique, which involves sitting on a branch or leaf and waiting for a female to fly by. Unlike other species of butterflies, such as the black swallowtail butterfly, P. polytes males neither defend territories nor lek, that is, gather to participate in a competitive display to attract a mate.

Males exhibit puddling behavior, which involves drinking from puddles, stream edges, coastlines, carrion, or feces. Puddling provides males with sodium and ammonia, which they pass to females via spermatophores. The intake of sodium and ammonia increases the reproductive success of males, female fecundity, and egg hatching success.

It is unknown whether P. polytes mates with multiple partners, but many butterfly species are promiscuous. Papilio polytes breeds throughout the year, but mostly in January through April. (Durgesh, et al., 2011; Honda, et al., 2012; Lederhouse, 1982; Pola and Garcia-Paris, 2005)

Papilio polytes breeds throughout the year, mostly in January through April. A male transfers a spermatophore (packet of sperm) to his mate. Females of most butterfly species store the sperm in an abdominal pouch called a spermatheca. When the female has a full pouch of sperm, she selects a host plant on which to lay her eggs. A typical female P. polytes places eggs on the upper and lower surfaces of a given leaf and lays hundreds of eggs in her lifetime. (Sehgal, et al., 2009; Shobana, et al., 2010; Spitzer, 1983; Suwarno, et al., 2010b)

  • Breeding season
    Throughout the year, mostly in January through April
  • Range eggs per season
    100 to 350

Male common mormon butterflies engage in puddling behavior to acquire sodium, ammonia, and other minerals from soil, puddles, seawater, carrion, and feces. During copulation, these minerals are transferred to a female through the spermatophore (packet containing sperm). The female passes these minerals into her eggs. This puddling behavior and the transfer of minerals can increase the reproductive success of adults and increase the survival rate of eggs.

Larval growth, development, and longevity depend on the nutrients in the host plant. Papilio polytes females preferentially lay eggs on host plants of higher nutritional quality, such as Citrus reticulata and Citrus aurantifolia. (Inoue, 2006; Molleman, 2009; Pola and Garcia-Paris, 2005; Suwarno, et al., 2010a; Suwarno, et al., 2010b)

  • Parental Investment
  • pre-fertilization
    • provisioning
    • protecting
      • male
      • female
  • pre-hatching/birth
    • provisioning
      • male
      • female

Lifespan/Longevity

Adult Papilio polytes females typically live longer than adult males, as females live an average of 6 to 8 days, while males live an average of 3 to 4 days. It is possible that the production of mimetic patterns, although beneficial by reducing predation, can also reduce the lifespan of mimetic P. polytes. Non-mimetic females can live longer than males and mimetic females, but they have a higher risk of predation. Females tend to be attacked more frequently than males because their larger abdomen is a more rewarding meal for predators.

The entire life cycle of P. polytes, from the laying of an egg to the death of an adult, can be about 30 to 43 days. (Ohsaki, 2005; Shobana, et al., 2010; Watanabe, 1979)

  • Typical lifespan
    Status: wild
    30 to 43 days

Behavior

One of the most unique adaptations of female common mormons is their mimicry of unpalatable butterfly species, including Pachliopta aristolochiae (common rose butterfly) and Pachliopta hector (crimson rose butterfly). Because the common mormon actually is edible by predators, and it mimics unpalatable species, the deception is an example of Batesian mimicry. Mimetic Papilio polytes females mimic not only the appearance, but also the flight pattern of P. aristolochiae; for example, the P. polytes polymorphism that mimics P. aristolochiae beats its wings slowly, with the wings held almost parallel to the ground, to prominently display the warning coloration marks on its hindwings. In contrast, P. polytes males and females with non-mimetic markings follow a swift, erratic flight pattern to escape predation.

Male common mormons engage in puddling behavior, which involves males congregating on and drinking from moist earth, puddles, stream edges, coastlines, carrion, or feces. By puddling, males can obtain sodium, ammonia, and other minerals that can improve their own reproductive success, the fecundity of their female mates, and egg hatching success. Groups of male butterflies typically puddle in cool, shady areas, rather than out in the open, where the risk of predation likely is higher.

Because they are larger than males, female common mormons bask longer in the mornings before they can fly. Both males and females bask with their wings spread open and pressed flat against their resting place. They usually bask on vegetation near the forest floor. After sundown, they rest on plant material close to the ground. (Barua, et al., 2004; Honda, et al., 2012; Kitamura and Imafuku, 2010; Ohsaki, 2005; Rangana, 2010; Sehgal, et al., 2009)

Home Range

The home range of Papilio polytes is >1 km; within the range, individuals from different populations come in contact with one another. Females tend to remain in one area for one day, while males stay three to four days. Because common mormon butterflies rarely fly over the undergrowth, they move to new territories by coasting on wind currents. (Watanabe, 1979)

Communication and Perception

Because butterflies have compound eyes, they can see in many directions simultaneously. Their eyes consist of many individual hexagonal light sensors, each angled in a slightly different direction. Their eyes can detect light, color, and motion.

Papilio polytes females carefully choose the plants on which to lay their eggs by using chemosensilla, or elongated bristles, to "taste" the chemical components of the vegetation. To investigate a plant, a female butterfly drums her forelegs on a leaf to gain information about plant chemistry via her chemotactile sensory receptors. Males might use olfaction to detect bodies of water that contain salt and minerals by using their antennae.

Papilio polytes males produce a pheromone in their wings. Because the pheromone is not produced by females, this chemical may be used by P. polytes females to differentiate between potential sexual partners and males of other species. The function of the chemicals produced by P. polytes males, however, has not been shown definitively. (Brinckerhoff, 2006; Inoue, 2006; Omura and Honda, 2005; Pola and Garcia-Paris, 2005)

Food Habits

Papilio polytes larvae are folivores, eating only the leaves of the host plant. A P. polytes larva has a pair of hypognathous mandibles, in which the lower mandible is longer than upper mandible; these structures enable the caterpillar to cut leaves into smaller pieces. The caterpillar eats the leaves of plants such as the key lime, sweet orange, sweet lime, bael, mandarin orange, kaffir lime, curry tree, gin berry, and orange climber. The caterpillars prefer new, fully opened leaves. The first- and second-instar caterpillars eat only the edges of the leaves. The third, fourth, and fifth instars eat the entire leaf, avoiding the veins. Larvae prefer plants with smooth leaves that have a high nitrogen and water content. Because these plants are grown agriculturally, common mormon caterpillars can damage crop yields.

Papilio polytes adults are nectarivores, feeding on nectar from short herbs and shrubs that have elongated corollar tubes, such as Lantana, Jatropha, Ixora and Mussaenda in urban areas, and Asystasia, Peristrophe, and Jasminum in rural areas. (Rangana, 2010; Sehgal, et al., 2009; Suwarno, et al., 2010a; Suwarno, et al., 2010b)

  • Plant Foods
  • leaves
  • nectar

Predation

Mortality rates for Papilio polytes are highest during the egg stage. In the egg stage, P. polytes is most susceptible to spiders of the genus Oxyopes. Other egg predators include the red ant and the parasitoid Ooencyrtus papilioni.

Early larval instars are attacked by Oxyopes spiders, such as the common lynx spider and the elegant lynx spider. Later larval instars are susceptible to a variety of predators, including birds, mantises, oriental garden lizards, and the assassin bug Sycanus dichotomus. The bodies of P. polytes larvae are colored in such a way that they resemble bird droppings, which camouflages them against predation. Later instars of the caterpillar have a structure called an osmeterium, which can be everted from the prothorax when the larva is threatened and produces a noxious compound that wards off predators.

A unique adaptation of female common mormon butterflies is their Batesian mimicry of the unpalatable butterfly species Pachlipta aristolochiae and Pachliopta hector. Although P. polytes is palatable, its mimicry of unpalatable species causes predators to avoid it. The mimetic females imitate both the appearance and the flight patterns of the unpalatable butterflies. Common rose butterflies, for example, beat their wings slowly, holding their wings almost parallel to the ground to display the warning coloration on their hindwings. Female common mormon butterflies mimicking the common rose also fly in a slow, regular pattern to make visible their warning markings. In contrast, adult male common mormons and adult females with non-mimetic markings follow a swift, erratic flight pattern to escape predation. (Hori, 2003; Kitamura and Imafuku, 2010; Ohsaki, 2005; Rangana, 2010; Sehgal, et al., 2009; Smetacek, 2006; Suwarno, 2010; Tan, 2011; Vasait, 2002)

Ecosystem Roles

Common mormon butterflies are pollinators, herbivores, and prey. The parasitoid Ooencyrtus papilioni attacks the eggs of Papilio polytes. (Durgesh, et al., 2011; Kitamura and Imafuku, 2010; Reddi and Bai, 1984; Shobana, et al., 2010; Smetacek, 2006; Suwarno, et al., 2010a; Suwarno, et al., 2010b; Suwarno, 2010; Tan, 2011; Vasait, 2002)

  • Ecosystem Impact
  • pollinates
Species Used as Host
  • Rutaceae
Commensal/Parasitic Species
  • Ooencyrtus papilioni

Economic Importance for Humans: Positive

Common mormon butterflies are an excellent example of sex-limited polymorphic mimicry and are used as a model system in studies of Batesian mimicry. Common mormon butterflies also do well in captivity and can be found in many butterfly conservatories. (Kitamura and Imafuku, 2010; Ohsaki, 2005; Smetacek, 2006)

  • Positive Impacts
  • research and education

Economic Importance for Humans: Negative

Papilio polytes larvae feed on many species of citrus and other plants, many of which are farmed commercially. Their diet includes mandarin orange, sweet lime and kaffir lime plants. Although the damage inflicted by P. polytes is relatively less than that caused by other herbivore species, it nonetheless is a pest on citrus crops. (Shobana, et al., 2010; Suwarno, et al., 2010b)

  • Negative Impacts
  • crop pest

Conservation Status

Papilio polytes has no special status; it is commonly found throughout its range. (Rangana, 2010; Suwarno, et al., 2010a)

Other Comments

In the common mormon, the females are heterogametic; that is, females carry two different sex chromosomes. The females have a Z and a W chromosome, while males carry two Z chromosomes. This is a common characteristic of all butterflies. (Presgraves, 2002)

Contributors

Kathryn Halloran (author), University of Michigan-Ann Arbor, Elizabeth Wason (author, editor), Animal Diversity Web Staff.

Glossary

Palearctic

living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.

World Map

agricultural

living in landscapes dominated by human agriculture.

aposematic

having coloration that serves a protective function for the animal, usually used to refer to animals with colors that warn predators of their toxicity. For example: animals with bright red or yellow coloration are often toxic or distasteful.

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

chemical

uses smells or other chemicals to communicate

cryptic

having markings, coloration, shapes, or other features that cause an animal to be camouflaged in its natural environment; being difficult to see or otherwise detect.

delayed fertilization

a substantial delay (longer than the minimum time required for sperm to travel to the egg) takes place between copulation and fertilization, used to describe female sperm storage.

diurnal
  1. active during the day, 2. lasting for one day.
ectothermic

animals which must use heat acquired from the environment and behavioral adaptations to regulate body temperature

fertilization

union of egg and spermatozoan

folivore

an animal that mainly eats leaves.

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

herbivore

An animal that eats mainly plants or parts of plants.

heterothermic

having a body temperature that fluctuates with that of the immediate environment; having no mechanism or a poorly developed mechanism for regulating internal body temperature.

internal fertilization

fertilization takes place within the female's body

metamorphosis

A large change in the shape or structure of an animal that happens as the animal grows. In insects, "incomplete metamorphosis" is when young animals are similar to adults and change gradually into the adult form, and "complete metamorphosis" is when there is a profound change between larval and adult forms. Butterflies have complete metamorphosis, grasshoppers have incomplete metamorphosis.

mimicry

imitates a communication signal or appearance of another kind of organism

motile

having the capacity to move from one place to another.

native range

the area in which the animal is naturally found, the region in which it is endemic.

nectarivore

an animal that mainly eats nectar from flowers

oriental

found in the oriental region of the world. In other words, India and southeast Asia.

World Map

oviparous

reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.

pheromones

chemicals released into air or water that are detected by and responded to by other animals of the same species

polymorphic

"many forms." A species is polymorphic if its individuals can be divided into two or more easily recognized groups, based on structure, color, or other similar characteristics. The term only applies when the distinct groups can be found in the same area; graded or clinal variation throughout the range of a species (e.g. a north-to-south decrease in size) is not polymorphism. Polymorphic characteristics may be inherited because the differences have a genetic basis, or they may be the result of environmental influences. We do not consider sexual differences (i.e. sexual dimorphism), seasonal changes (e.g. change in fur color), or age-related changes to be polymorphic. Polymorphism in a local population can be an adaptation to prevent density-dependent predation, where predators preferentially prey on the most common morph.

rainforest

rainforests, both temperate and tropical, are dominated by trees often forming a closed canopy with little light reaching the ground. Epiphytes and climbing plants are also abundant. Precipitation is typically not limiting, but may be somewhat seasonal.

semelparous

offspring are all produced in a single group (litter, clutch, etc.), after which the parent usually dies. Semelparous organisms often only live through a single season/year (or other periodic change in conditions) but may live for many seasons. In both cases reproduction occurs as a single investment of energy in offspring, with no future chance for investment in reproduction.

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

sperm-storing

mature spermatozoa are stored by females following copulation. Male sperm storage also occurs, as sperm are retained in the male epididymes (in mammals) for a period that can, in some cases, extend over several weeks or more, but here we use the term to refer only to sperm storage by females.

suburban

living in residential areas on the outskirts of large cities or towns.

tactile

uses touch to communicate

terrestrial

Living on the ground.

tropical

the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.

tropical savanna and grassland

A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.

savanna

A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.

temperate grassland

A terrestrial biome found in temperate latitudes (>23.5? N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.

urban

living in cities and large towns, landscapes dominated by human structures and activity.

visual

uses sight to communicate

year-round breeding

breeding takes place throughout the year

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