Papilio Demodocus
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''Papilio demodocus'', the citrus swallowtail or Christmas butterfly, is a
swallowtail butterfly Swallowtail butterflies are large, colorful butterflies in the family Papilionidae, and include over 550 species. Though the majority are tropical, members of the family inhabit every continent except Antarctica. The family includes the large ...
which commonly occurs over the entirety of
sub-Saharan Africa Sub-Saharan Africa is, geographically, the area and regions of the continent of Africa that lies south of the Sahara. These include West Africa, East Africa, Central Africa, and Southern Africa. Geopolitically, in addition to the List of sov ...
, including
Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Africa ...
, besides the southern
Arabian Peninsula The Arabian Peninsula, (; ar, شِبْهُ الْجَزِيرَةِ الْعَرَبِيَّة, , "Arabian Peninsula" or , , "Island of the Arabs") or Arabia, is a peninsula of Western Asia, situated northeast of Africa on the Arabian Plate ...
. The
caterpillars Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths). As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder Symph ...
feed on various native plants of especially the family
Rutaceae The Rutaceae is a family, commonly known as the rueRUTACEAE
in BoDD – Botanical Derm ...
, but have also taken to the leaves of cultivated
citrus ''Citrus'' is a genus of flowering plant, flowering trees and shrubs in the rue family, Rutaceae. Plants in the genus produce citrus fruits, including important crops such as Orange (fruit), oranges, Lemon, lemons, grapefruits, pomelos, and lim ...
trees.


Life cycle

Citrus swallowtails pass through approximately three generations per year.


Eggs

Female butterflies lay their eggs singly on citrus leaves. After about six days, the egg hatches into an immature
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
.


Immature larva

The immature larvae are black, yellow, and white with spikes. Their coloration provides effective
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the ...
, as they resemble bird droppings. They grow to a length of 10 or 15 mm before changing into mature larvae.


Mature larva

Mature larvae are green with white or pink markings and eyespots. They grow to a maximum length of about 45 mm. Mature
caterpillars Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths). As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder Symph ...
lack the camouflage of their immature state. Instead, when threatened by a
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...
or other predator, they produce a forked, orange-coloured organ known as an
osmeterium The osmeterium is a defensive organ found in all papilionid larvae, in all stages. The organ is situated in the prothoracic segment and can be everted when the larva feels threatened. The everted organ resembles a fleshy forked tongue (not unlik ...
. The organ emits a strong smell which acts as a discouragement to the predator. The larvae are sometimes known as 'orange dogs'.


Pupa

The caterpillars attach themselves to branches with
silk Silk is a natural protein fiber, some forms of which can be woven into textiles. The protein fiber of silk is composed mainly of fibroin and is produced by certain insect larvae to form cocoons. The best-known silk is obtained from the coc ...
, transforming into
pupa A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their ...
e. They remain in the pupal form for two to three weeks before emerging as adults.


Adult

Adult butterflies have black and yellow markings with red and blue eyespots. Female butterflies tend to be larger than males.


Natural enemies

As with most butterflies, various
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
s,
parasitoid In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionarily stable str ...
s and diseases attack ''Papilio demodocus'', so that
integrated pest management Integrated pest management (IPM), also known as integrated pest control (IPC) is a broad-based approach that integrates both chemical and non-chemical practices for economic control of pests. IPM aims to suppress pest populations below the econ ...
is generally the most rational approach to control of infestations, paying due attention to avoiding destruction of the populations of enemies. In particular
parasitic wasp Parasitoid wasps are a large group of hymenopteran superfamilies, with all but the wood wasps (Orussoidea) being in the wasp-waisted Apocrita. As parasitoids, they lay their eggs on or in the bodies of other arthropods, sooner or later causin ...
s in the family
Encyrtidae Encyrtidae is a large family of parasitic wasps, with some 3710 described species in about 455 genera. The larvae of the majority are primary parasitoids on Hemiptera, though other hosts are attacked, and details of the life history can be variab ...
, such as some species of the
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
''
Ooencyrtus ''Ooencyrtus'' is a genus of chalcid wasp. William Harris Ashmead named and circumscribed In geometry, the circumscribed circle or circumcircle of a polygon is a circle that passes through all the vertices of the polygon. The center of this ...
'' develop in ''Papilio'' eggs. Other parasitoidal wasps in families such as
Chalcididae The Chalcididae are a moderate-sized family within the Chalcidoidea, composed mostly of parasitoids and a few hyperparasitoids. The family is apparently polyphyletic, though the different subfamilies may each be monophyletic, and some may be elev ...
and
Braconidae The Braconidae are a family of parasitoid wasps. After the closely related Ichneumonidae, braconids make up the second-largest family in the order Hymenoptera, with about 17,000 recognized species and many thousands more undescribed. One analysis ...
(for example genus ''
Apanteles ''Apanteles'' is a very large genus of Braconidae, braconid wasps, containing more than 600 described species found worldwide. There are no native species in New Zealand, and none have been recorded in the high arctic. See also * List of Apantel ...
'') attack the larvae, and
Pteromalidae The Pteromalidae are a very large family of mostly parasitoid wasps, with some 3,450 described species in about 640 genera (the number was greater, but many species and genera have been reduced by synonymy in recent years). The subfamily-level d ...
(for example genus '' Pteromalus'') are parasitoids of the pupae. Predatory insects such as certain
Heteroptera The Heteroptera are a group of about 40,000 species of insects in the order Hemiptera. They are sometimes called "true bugs", though that name more commonly refers to the Hemiptera as a whole. "Typical bugs" might be used as a more unequivocal al ...
, in particular
Reduviidae The Reduviidae are a large cosmopolitan family of the order Hemiptera (true bugs). Among the Hemiptera and together with the Nabidae almost all species are terrestrial ambush predators: most other predatory Hemiptera are aquatic. The main exampl ...
known as assassin bugs, and some
Pentatomidae Pentatomidae is a family of insects belonging to the order Hemiptera, generally called shield bugs or stink bugs. Pentatomidae is the largest family in the superfamily Pentatomoidea, and contains around 900 genera and over 4700 species.Robert G ...
attack the larvae, and certain
Mantodea Mantises are an order (Mantodea) of insects that contains over 2,400 species in about 460 genera in 33 families. The largest family is the Mantidae ("mantids"). Mantises are distributed worldwide in temperate and tropical habitats. They ha ...
such as genus '' Sphodromantis'' attack both larvae and adults.


Taxonomy

''Papilio demodocus'' is a member of the ''demoleus''
species group In biology, a species complex is a group of closely related organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The taxa in the complex may be able to hybridize readily with each oth ...
. The
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
members are: *''Papilio demodocus'' Esper, 1799 *'' Papilio demoleus'' Linnaeus, 1758 *'' Papilio erithonioides'' Grose-Smith, 1891 *'' Papilio grosesmithi'' Rothschild, 1926 *''
Papilio morondavana ''Papilio morondavana'', the Madagascan emperor swallowtail, is a species of butterfly in the family Papilionidae. It is endemic to Madagascar. The habitat consists of forests. Original description Expanse male 4 and 3/8, female 4 and 3/4 inche ...
'' Grose-Smith, 1891


Subspecies

*''Papilio demodocus demodocus'' —
sub-Saharan Africa Sub-Saharan Africa is, geographically, the area and regions of the continent of Africa that lies south of the Sahara. These include West Africa, East Africa, Central Africa, and Southern Africa. Geopolitically, in addition to the List of sov ...
, including
Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Africa ...
*''Papilio demodocus bennetti'' Dixey, 1898
Socotra Socotra or Soqotra (; ar, سُقُطْرَىٰ ; so, Suqadara) is an island of the Republic of Yemen in the Indian Ocean, under the ''de facto'' control of the UAE-backed Southern Transitional Council, a secessionist participant in Yemen’s ...


Mating system


Courtship

Courtship is more or less the same in all Papilionidae. Once a female enters the visual field of a male, the male moves quickly to hover over her so that his wings beat rapidly. The female is then induced to land so that the male can attempt to mate with her. There are various ways in which the male entices the female, including visual, olfactory, tactile, and auditory cues. Of particular interest is the use of olfactory cues. Male butterflies produce
pheromones A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
from different structures, such as that of the anal fold of the hindwing, that cause the females to perform the appropriate response. Sometimes, however, females choose to reject a male's attempt at mating, often because she has already mated. She can do so by either avoiding his approach or, if she lands, she will flap her wings quickly and deliberately all while raising her abdomen until the male flies away. In this way, courtship is primarily a female's choice.Douglas, Matthew M. ''The Lives of Butterflies''. Ann Arbor: University of Michigan, 1986. Print. Furthermore, it has been observed that females also produce a pheromone that aids males in determining whether a female has already mated or not.


Mating

This species primarily mates via the
lek Lek or LEK may refer to: * Lek mating, mating in a lek, a type of animal territory in which males of a species gather * Albanian lek, the currency of Albania * Lek (magazine), a Norwegian softcore pornographic magazine * Lek (pharmaceutical comp ...
system, in which there are aggregations of males on small mating territories. When the female reaches the lek, she changes her behavior so that she helps the males to detect her by performing a long and obvious circular flight. The species operates on a
polygynous Polygyny (; from Neoclassical Greek πολυγυνία (); ) is the most common and accepted form of polygamy around the world, entailing the marriage of a man with several women. Incidence Polygyny is more widespread in Africa than in any ...
system in which one male has the ability to mate with several females in one breeding season.Boggs, Carol L., Ward B. Watt, and Paul R. Ehrlich. ''Butterflies: Ecology and Evolution Taking Flight''. Chicago: University of Chicago, 2003. Print. In ''
Papilio glaucus ''Papilio glaucus'', the eastern tiger swallowtail, is a species of butterfly native to eastern North America. It is one of the most familiar butterflies in the eastern United States, ranging north to southern Ontario, Canada, and is common in m ...
'', that which is of the same genus of this species, the lack of male-male competition, strong rapid flight, dispersed abundant food, and
oviposition The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typical ...
sights helped to support the idea of mating system based on polygyny. The females are prevented from mating with other males when the male emits a sphragis, which prevents other males from mating with the female, ensuring that only the sperm from this male fertilizes the eggs. The physical act of copulation takes between one half hour to two hours. During this time, the male
spermatophore A spermatophore or sperm ampulla is a capsule or mass containing spermatozoa created by males of various animal species, especially salamanders and arthropods, and transferred in entirety to the female's ovipore during reproduction. Spermatophores ...
is transferred to the female's bursa. The spermatophore is then absorbed and its nutrient (protein) contents are given to the female's eggs as a food source. For this reason, the male makes an indirect investment to his offspring, given that he offers them a food source, being beneficial to the female given that she risks less of her own well-being to bear her offspring. Therefore, a male is considered more fit with a larger ejaculation, given that he allows his offspring to feed so that his own genes may be passed on efficiently.


Mistakes and diversity

Mate assessment involves the discrimination of
conspecifics Biological specificity is the tendency of a characteristic such as a behavior or a biochemical variation to occur in a particular species. Biochemist Linus Pauling stated that "Biological specificity is the set of characteristics of living organis ...
, so that mating requires that males and females recognize and mate with individuals of the appropriate sex and species, as indicated by the appropriate chemical and visual cues that are made. Sometimes, homosexual copulation has been seen in addition to two males upon one female. Very rarely do females accept a mate from a different species or genus, and if they do, it is usually because they are too immature to realize the correct sexual cues of males from their own species. However, these "incorrect" matings do occur so that there is diversity in the species, as seen through the different larval patterns that have evolved in the species, depending on the type of plant the eggs are laid upon.Clarke, C. A., C.G.C. Dickson, and P.M. Sheppard. "Larval Color Pattern in ''Papilio demodocus''." Evolution 17.2 (June 1963): 130-37. JSTOR. Web.17 Sept. 2013.


Gallery

File:Citrus swallowtail larva.JPG, An ectoparasitic fly, ''cf.''
Ceratopogonidae Ceratopogonidae is a family of flies commonly known as no-see-ums, or biting midges, generally in length. The family includes more than 5,000 species, distributed worldwide, apart from the Antarctic and the Arctic. Ceratopogonidae are holomet ...
, sucking blood from a larva of ''P. demodocus'' Papilio demodocus larva IMG 8543s.jpg, Final instar larva,
osmeterium The osmeterium is a defensive organ found in all papilionid larvae, in all stages. The organ is situated in the prothoracic segment and can be everted when the larva feels threatened. The everted organ resembles a fleshy forked tongue (not unlik ...
extended, applying repugnatorial secretion with one "horn" to the hand holding it. File:Papilio demodocus pupa 2.jpg, Mid-stage ''Papilio demodocus'' pupa File:Papilio demodocus pupa 3.jpg, Pupa near
eclosion A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their ...
, the colour pattern of adult visible through the skin


References

{{Taxonbar, from=Q133397 demodocus Agricultural pest insects Fauna of Rivers State Butterflies of Africa Butterflies described in 1798 Lepidoptera of Cape Verde