Citrus Swallowtail (Papilio Demodocus) On Elephant Dung
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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 larg ...
which commonly occurs over the entirety of
sub-Saharan Africa Sub-Saharan Africa is the area and regions of the continent of Africa that lie south of the Sahara. These include Central Africa, East Africa, Southern Africa, and West Africa. Geopolitically, in addition to the list of sovereign states and ...
, including
Madagascar Madagascar, officially the Republic of Madagascar, is an island country that includes the island of Madagascar and numerous smaller peripheral islands. Lying off the southeastern coast of Africa, it is the world's List of islands by area, f ...
, besides the southern
Arabian Peninsula The Arabian Peninsula (, , or , , ) or Arabia, is a peninsula in West Asia, situated north-east of Africa on the Arabian plate. At , comparable in size to India, the Arabian Peninsula is the largest peninsula in the world. Geographically, the ...
. 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 Sym ...
feed on various native plants of especially the family
Rutaceae The Rutaceae () is a family (biology), family, commonly known as the rueRUTACEAE
in Bo ...
, but have also taken to the leaves of cultivated
citrus ''Citrus'' is a genus of flowering trees and shrubs in the family Rutaceae. Plants in the genus produce citrus fruits, including important crops such as oranges, mandarins, lemons, grapefruits, pomelos, and limes. ''Citrus'' is nativ ...
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 (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
.


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 b ...
, 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 Sym ...
lack the camouflage of their immature state. Instead, when threatened by a
bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
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 fiber, natural protein fiber, some forms of which can be weaving, woven into textiles. The protein fiber of silk is composed mainly of fibroin and is most commonly produced by certain insect larvae to form cocoon (silk), c ...
, transforming into
pupa A pupa (; : 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 life cycle, the stages th ...
e. They remain in the pupal form for two to three weeks before emerging as adults.It is dormint for four months in the pupa from May to August. Pupa will amerge in September


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 in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
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) integrates both chemical and non-chemical practices for economic control of pests. The UN's Food and Agriculture Organization defines IPM as "the careful consideratio ...
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 causi ...
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 (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
'' Ooencyrtus'' 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 e ...
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 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 Apanteles species ...
'') attack the larvae, and
Pteromalidae The Pteromalidae are a large family of wasps, the majority being parasitoids of other insects. They are found throughout the world in virtually all habitats, and many are important as biological control agents. The oldest known fossil is known fr ...
(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 is a large Cosmopolitan distribution, cosmopolitan family of the suborder Heteroptera of the Order (biology), order Hemiptera (true bugs). Among the Hemiptera and together with the Nabidae almost all species are terrestrial ambush ...
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 (biology), order (Mantodea) of insects that contains List of mantis genera and species, over 2,400 species in about 460 genera in 33 families. The largest family is the Mantidae ("mantids") ...
such as genus ''
Sphodromantis ''Sphodromantis'' is a large genus of praying mantises concentrated in Africa, sometimes considered a synonym of the genus ''Hierodula'': from the same family, Mantidae. Outside their range especially, many share the common name African Mantis. ...
'' 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 In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
members are: *''Papilio demodocus'' Esper, 1799 *'' Papilio demoleus'' Linnaeus, 1758 *'' Papilio erithonioides'' Grose-Smith, 1891 *'' Papilio grosesmithi'' Rothschild, 1926 *'' Papilio morondavana'' Grose-Smith, 1891


Subspecies

*''Papilio demodocus demodocus'' —
sub-Saharan Africa Sub-Saharan Africa is the area and regions of the continent of Africa that lie south of the Sahara. These include Central Africa, East Africa, Southern Africa, and West Africa. Geopolitically, in addition to the list of sovereign states and ...
, including
Madagascar Madagascar, officially the Republic of Madagascar, is an island country that includes the island of Madagascar and numerous smaller peripheral islands. Lying off the southeastern coast of Africa, it is the world's List of islands by area, f ...
*''Papilio demodocus bennetti'' Dixey, 1898
Socotra Socotra, locally known as Saqatri, is a Yemeni island in the Indian Ocean. Situated between the Guardafui Channel and the Arabian Sea, it lies near major shipping routes. Socotra is the largest of the six islands in the Socotra archipelago as ...


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 behavi ...
from different structures, such as that of the anal fold of the hindwing, that cause the females to perform the appropriate response. Sometimes, however, a female can 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 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 () is a form of polygamy entailing the marriage of a man to several women. The term polygyny is from Neoclassical Greek πολυγυνία (); . Incidence Polygyny is more widespread in Africa than in any other continent. Some scholar ...
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 ma ...
'', 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 typica ...
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, from Ancient Greek σπέρμα (''spérma''), meaning "seed", and -φόρος (''-phóros''), meaning "bearing", or sperm ampulla is a capsule or mass containing spermatozoa created by males of various animal species, especiall ...
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 organism ...
, 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

Papilio demodocus larva 0.jpg, Early
instar An instar (, from the Latin '' īnstar'' 'form, likeness') is a developmental stage of arthropods, such as insects, which occurs between each moult (''ecdysis'') until sexual maturity is reached. Arthropods must shed the exoskeleton in order to ...
File:Citrus swallowtail larva.JPG, An ectoparasitic fly, ''cf.''
Ceratopogonidae Ceratopogonidae is a family of flies commonly known as no-see-ums, sand flies or biting midges, generally in length. The family includes more than 5,000 species, distributed worldwide, apart from the Antarctic and the Arctic. A 2025 study fro ...
, sucking blood from a larva 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 pupa File:Papilio demodocus pupa 3.jpg, Pupa near
eclosion A pupa (; : 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 life cycle, the stages th ...
, the colour pattern of adult visible through the skin Citrus swallowtails (Papilio demodocus) Principe.jpg, a pair nectaring on hibiscus


References

{{Taxonbar, from=Q133397 demodocus Agricultural pest insects Fauna of Rivers State Butterflies of Africa Butterflies described in 1798 Lepidoptera of Cape Verde Taxa named by Eugenius Johann Christoph Esper Butterflies of Asia