''Danaus chrysippus'', also known as the plain tiger, African queen, or African monarch, is a medium-sized butterfly widespread in

Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an are ...

, Australia and

Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...

. It belongs to the

Danainae Danainae is a subfamily of the family Nymphalidae, the brush-footed butterflies. It includes the Daniadae, or milkweed butterflies, who lay their eggs on various milkweeds on which their larvae (caterpillars) feed, as well as the clearwing butt ...

subfamily of the brush-footed butterfly

family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...

Nymphalidae The Nymphalidae are the largest family of butterflies, with more than 6,000 species distributed throughout most of the world. Belonging to the superfamily Papilionoidea, they are usually medium-sized to large butterflies. Most species have a red ...

. Danainae primarily consume plants in the genus ''

Asclepias ''Asclepias'' is a genus of herbaceous, perennial, flowering plants known as milkweeds, named for their latex, a milky substance containing cardiac glycosides termed cardenolides, exuded where cells are damaged. Most species are toxic to humans ...

'', more commonly called milkweed. Milkweed contains toxic compounds,

cardenolide A cardenolide is a type of steroid. Many plants contain derivatives, collectively known as cardenolides, including many in the form of cardenolide glycosides (cardenolides that contain structural groups derived from sugars). Cardenolide glycoside ...

s, which are often consumed and stored by many butterflies. Because of their

emetic Vomiting (also known as emesis and throwing up) is the involuntary, forceful expulsion of the contents of one's stomach through the mouth and sometimes the nose. Vomiting can be the result of ailments like food poisoning, gastroenteritis ...

properties, the plain tiger is

unpalatable Palatability (or palatableness) is the hedonic reward (i.e., pleasure) provided by foods or fluids that are agreeable to the "palate", which often varies relative to the homeostatic satisfaction of nutritional, water, or energy needs. The palatabi ...

to most predators. As a result, its coloration is widely mimicked by other

species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...

of butterflies. The plain tiger inhabits a wide variety of

habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...

s, although it is less likely to thrive in

jungle A jungle is land covered with dense forest and tangled vegetation, usually in tropical climates. Application of the term has varied greatly during the past recent century. Etymology The word ''jungle'' originates from the Sanskrit word ''ja� ...

-like conditions and is most often found in drier, wide-open areas. ''D. chrysippus'' encompasses three main subspecies: ''D. c. alcippus, D. c. chrysippus,'' and ''D. c. orientis.'' These subspecies are found concentrated in specific regions within the larger range of the entire species. The plain tiger is believed to be one of the first butterflies depicted in art. A 3,500-year-old

ancient Ancient history is a time period from the beginning of writing and recorded human history to as far as late antiquity. The span of recorded history is roughly 5,000 years, beginning with the Sumerian cuneiform script. Ancient history cov ...

Egypt Egypt ( ar, مصر , ), officially the Arab Republic of Egypt, is a transcontinental country spanning the northeast corner of Africa and southwest corner of Asia via a land bridge formed by the Sinai Peninsula. It is bordered by the Medit ...

ian fresco in

Luxor Luxor ( ar, الأقصر, al-ʾuqṣur, lit=the palaces) is a modern city in Upper (southern) Egypt which includes the site of the Ancient Egyptian city of ''Thebes''. Luxor has frequently been characterized as the "world's greatest open-a ...

features the oldest known illustration of this species.

Description

''D. chrysippus'' is a medium-sized

butterfly Butterflies are insects in the macrolepidopteran clade Rhopalocera from the order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The group comprise ...

with a

wingspan The wingspan (or just span) of a bird or an airplane is the distance from one wingtip to the other wingtip. For example, the Boeing 777–200 has a wingspan of , and a wandering albatross (''Diomedea exulans'') caught in 1965 had a wingspan o ...

of about . The body is black with many white spots. The wings are orange, the upperside brighter and richer than the underside. The apical half of the forewing is black with a white band. The hindwing has three black spots in the center. The wings are bordered in black and outlined with semicircular white spots. This species exhibits slight sexual dimorphism, as the Male has large scent glands on his hindwings, which the female lacks. They appear as a large black spot with a white centre if viewed from the underside ''D. chrysippus'' is a polymorphic species, so the exact coloring and patterning vary within and between

population Population typically refers to the number of people in a single area, whether it be a city or town, region, country, continent, or the world. Governments typically quantify the size of the resident population within their jurisdiction using a ...

s. It is similar in appearance to the Indian fritillary ('' Argynnis hyperbius''), which may coexist with it. File:Plain tiger butterfly wing scales.jpg, Close-up of the wing scales of a male ''Danaus chrysippus'' File:Danaus chrysippus male 2 by kadavoor.jpg, Male showing the pheromone pouch and brush-like organ in Kerala

Geographic range

The plain tiger is found across the entirety of Africa, where the predominant subspecies is ''D. c. alcippus''. Its range extends across the majority of Asia throughout

Indian subcontinent The Indian subcontinent is a list of the physiographic regions of the world, physiographical region in United Nations geoscheme for Asia#Southern Asia, Southern Asia. It is situated on the Indian Plate, projecting southwards into the Indian O ...

, as well as many south

Pacific The Pacific Ocean is the largest and deepest of Earth's five oceanic divisions. It extends from the Arctic Ocean in the north to the Southern Ocean (or, depending on definition, to Antarctica) in the south, and is bounded by the contine ...

islands. The plain tiger is even present in parts of Australia. ''D. c. chrysippus'' is most common throughout Asia and in some select regions in Africa, while ''D. c. orientis'' is present in more

tropical The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...

African regions as well as some African islands, 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 ...

and the Seychelles. It is also found in Southern Europe and Kuwait. These insects are considered bioinvaders in North America.

Habitat

The plain tiger prefers

arid A region is arid when it severely lacks available water, to the extent of hindering or preventing the growth and development of plant and animal life. Regions with arid climates tend to lack vegetation and are called xeric or desertic. Most ...

, open areas, and is found in a variety of habitats, including deserts,

mountain A mountain is an elevated portion of the Earth's crust, generally with steep sides that show significant exposed bedrock. Although definitions vary, a mountain may differ from a plateau in having a limited summit area, and is usually highe ...

deciduous In the fields of horticulture and Botany, the term ''deciduous'' () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed leaves, usually in the autumn; to the shedding of petals, ...

forests, and human-tended

garden A garden is a planned space, usually outdoors, set aside for the cultivation, display, and enjoyment of plants and other forms of nature. The single feature identifying even the wildest wild garden is ''control''. The garden can incorporate bot ...

s in cities and parks. It is comfortable at altitudes ranging from sea level to around .

Food resources

Larval food plants

The plain tiger's larval host plants are from several families, most importantly

Asclepiadoideae The Asclepiadoideae are a subfamily of plants in the family Apocynaceae. Formerly, they were treated as a separate family under the name Asclepiadaceae, e.g. by APG II, and known as the milkweed family. They form a group of perennial herbs, twi ...

(

Apocynaceae Apocynaceae (from ''Apocynum'', Greek for "dog-away") is a family of flowering plants that includes trees, shrubs, herbs, stem succulents, and vines, commonly known as the dogbane family, because some taxa were used as dog poison Members of ...

): * ''

'' – milkweeds (recorded on ''A. cancellata'', ''A. coarctata'', '' A. curassavica'', ''A. fulva'', ''A. kaessneri'', ''A. lineolata'', '' A. physocarpa'', ''A. reflexa'', ''A. scabrifolia'', ''A. semilunata'', ''A. stenophylla'', ''A. swynnertonii'', '' A.syriaca'') * ''Aspidoglossum interruptum'' * '' Calotropis'' – mudar (recorded on '' C. gigantea'', '' C. procera'') * '' Caralluma burchardii'' (recorded from Canary Islands/Spain) * '' Ceropegia dichotoma'' (recorded from Canary Islands/Spain) * '' Cryptolepis buchananii'' * ''

Cynanchum ''Cynanchum'' is a genus of about 300 species including some swallowworts, belonging to the family Apocynaceae. The taxon name comes from Greek ''kynos'' (meaning "dog") and ''anchein'' ("to choke"), hence the common name for several species is d ...

'' (recorded on ''C. abyssinicum'', '' C. acutum'', ''C. altiscandens'', ''C. amplexicaule'', ''C. carnosum'', ''C. floribundum'', ''C. sublanceolatum'') * '' Gomphocarpus fruticosus'' * ''Kanahia laniflora'' * '' Leichardtia australis'' * ''Leptadenia hastata'' * '' Marsdenia leichhardtiana'' * ''Metaplexis japonica'' * '' Orbea variegata'' (recorded from Canary Islands/Spain) * ''Oxystelma pulchellum'' * '' Pentatropis'' (recorded on ''P. atropurpurea'', ''P. quinquepartita'') * ''

Pergularia daemia ''Pergularia daemia'', the trellis-vine, is a hispid, perennial vine in the family Asclepiadaceae, with an extensive range in the Old World tropics and subtropics. It has been used traditionally to treat a number of ailments. It is sometimes call ...

'' * ''Periploca linearifolia'' * ''Pleurostelma cernuum'' * '' Secamone'' (recorded on ''S. afzelii'', ''S. parvifolia'', ''S. platystigma'') * '' Stapelia gigantea'' * '' Stathmostelma'' (recorded on ''S. gigantiflorum'', ''S. pedunculatum'') * ''

Tylophora ''Tylophora'' is a genus of climbing plant or vine, first described as a genus in 1810. It is native to tropical and subtropical Asia, Africa, and Australia. Most of the species are perennial lianas. The name is derived from the Ancient Greek '' ...

'' (recorded on ''T. stenoloba'', ''T. sylvatica'') Host plants from other families include ''Dyerophytum indicum'' (

Plumbaginaceae Plumbaginaceae is a family of flowering plants, with a cosmopolitan distribution. The family is sometimes referred to as the leadwort family or the plumbago family. Most species in this family are perennial herbaceous plants, but a few grow ...

), ''

Ficus ''Ficus'' ( or ) is a genus of about 850 species of woody trees, shrubs, vines, epiphytes and hemiepiphytes in the family Moraceae. Collectively known as fig trees or figs, they are native throughout the tropics with a few species extendi ...

'' (

Moraceae The Moraceae — often called the mulberry family or fig family — are a family of flowering plants comprising about 38 genera and over 1100 species. Most are widespread in tropical and subtropical regions, less so in temperate climates; however ...

; recorded on ''F. laevis'', '' F. racemosa''), ''

Ipomoea ''Ipomoea'' () is the largest genus in the plant family Convolvulaceae, with over 600 species. It is a large and diverse group, with common names including morning glory, water convolvulus or water spinach, sweet potato, bindweed, moonfl ...

'' (

Convolvulaceae Convolvulaceae (), commonly called the bindweeds or morning glories, is a family of about 60 genera and more than 1,650 species. These species are primarily herbaceous vines, but also include trees, shrubs and herbs. The tubers of several spe ...

; recorded on '' I. alba'', '' I. bona-nox''), ''Lepisanthes rubiginosa'' ( Sapindaceae) as well as some

Euphorbiaceae Euphorbiaceae, the spurge family, is a large family of flowering plants. In English, they are also commonly called euphorbias, which is also the name of a genus in the family. Most spurges, such as '' Euphorbia paralias'', are herbs, but some, ...

Malvaceae Malvaceae, or the mallows, is a family of flowering plants estimated to contain 244 genera with 4225 known species. Well-known members of economic importance include okra, cotton, cacao and durian. There are also some genera containing familiar ...

, Poaceae, Rosaceae and

Scrophulariaceae The Scrophulariaceae are a family of flowering plants, commonly known as the figwort family. The plants are annual and perennial herbs, as well as shrubs. Flowers have bilateral (zygomorphic) or rarely radial (actinomorphic) symmetry. The Scr ...

Adult food plants

Adult plain tiger butterflies obtain nectar from various

flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...

s. The particular plants available vary depending on the geographic range of the butterfly population and the season, as certain plants do not flower throughout the entire year.

India

* ''

Catharanthus roseus ''Catharanthus roseus'', commonly known as bright eyes, Cape periwinkle, graveyard plant, Madagascar periwinkle, old maid, pink periwinkle, rose periwinkle, is a species of flowering plant in the family Apocynaceae. It is native and endemic to M ...

'' * ''

Lantana camara ''Lantana camara'' (common lantana) is a species of flowering plant within the verbena family ( Verbenaceae), native to the American tropics. It is a very adaptable species, which can inhabit a wide variety of ecosystems; once it has been introdu ...

'' * '' Vernonia cinerea'' * '' Tridax procumbens'' * '' Asystasia gangetica'' * ''

Antigonon leptopus ''Antigonon leptopus'' is a species of perennial vine in the buckwheat family commonly known as coral vine or queen's wreath. This clambering vine is characterized by showy, usually pink flowers that can bloom throughout the year and large, heart ...

'', January through April and August through December * ''

Tecoma stans ''Tecoma stans'' is a species of flowering perennial shrub in the trumpet vine family, Bignoniaceae, that is native to the Americas. Common names include yellow trumpetbush, yellow bells, yellow elder, ginger-thomas. ''Tecoma stans'' is the o ...

'', May through December * '' Heliotropium indicum''

Australia

* ''Goodenia maidenian'' * '' Eucalyptus conglobata'' * ''E. oleosa'' * ''Ptilotis obovata'' * ''

'' * ''

Lantana ''Lantana'' () is a genus of about 150 species of perennial flowering plants in the verbena family, Verbenaceae. They are native to tropical regions of the Americas and Africa but exist as an introduced species in numerous areas, especially in ...

'' * '' Leucopogon'' * ''

Daviesia ''Daviesia'', commonly known as bitter-peas, is a genus of about 130 species of flowering plants in the family Fabaceae, and is endemic to Australia. Plants in the genus ''Daviesia'' are shrubs or small trees with leaves modified as phyllodes or ...

'' In addition to nectar, adult plain tigers obtain pyrrolizidine alkaloids from the dead stems of different plant types. In Australia, the following plants have been identified as sources of pyrrolizidine alkaloids for ''D. chrysippus:'' * ''Parsonsia eucalyptophylla'' * ''P. straminea'' * '' Echium plantogineum'' * ''Senecio pterophorus'' * '' Heliotropium amplexicaule''

Parental care

Oviposition

Females lay eggs singly on the underside of the leaves of a larval food plant. The eggs are most often laid close to the ground.

Life history

Egg

The egg of the plain tiger is about long and across. When first laid it is white, but gradually turns brown over time. The egg is ridged and dome-shaped. Depending on temperature, the egg is typically hatched in 3–5 days. File:Plain tiger butterfly egg.jpg, Egg at

Tamil Nadu Tamil Nadu (; , TN) is a state in southern India. It is the tenth largest Indian state by area and the sixth largest by population. Its capital and largest city is Chennai. Tamil Nadu is the home of the Tamil people, whose Tamil language ...

, India File:Plain tiger egg.JPG, At Mananpur,

Mumbai Mumbai (, ; also known as Bombay — List of renamed Indian cities and states#Maharashtra, the official name until 1995) is the capital city of the Indian States and union territories of India, state of Maharashtra and the ''de facto'' fin ...

, Maharashtra

Caterpillar

The

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. ...

e of ''D. chrysippus'' proceeds through five

instar An instar (, from the Latin '' īnstar'', "form", "likeness") is a developmental stage of arthropods, such as insects, between each moult (''ecdysis''), until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or ...

stages. The first instar is about long, and its body is white while the head is black. The second instar is about long, and its body is primarily gray with yellow and black horizontal stripes. This coloration remains for the final three instar stages. The third instar is about long, the fourth about long, and the fifth about long. Depending on temperature, the larval stage can last from 12 to 20 days. File:Plain tiger larvae just hatched and eat its own eggshell..JPG, Larva eating its own eggshell File:Plain tiger moat.JPG, Caterpillar making " moat" to feed during the first instar File:Danaus chrysippus second instar.JPG, Second instar File:Plain Tiger Danaus chrysippus Caterpillar on Calotropis procera 2. by Dr. Raju Kasambe.jpg, Mature caterpillar File:Plain Tiger (Danaus chrysippus) caterpillar on a Calotropis (Milkweed) species in Hyderabad, AP W IMG 7970.jpg, Older caterpillar

Prepupal and pupal stages

Before

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 thei ...

tion, the caterpillar will become motionless and cease feeding. Its color shifts slightly from gray to brown, and it may lose a small amount of body mass. The prepupal stage lasts 1–3 days depending on temperature. The pupal stage lasts 9–15 days depending on temperature, and the pupa changes color over this period from a pale green to dark brown. Pupae are about tall and wide. File:Chrysalide Danaus chrysippus.jpg, Pupa File:Plain tiger pupa.jpg, Fresh pupa at Bandra Kurla Complex, Mumbai File:Danauschrysippus pupa sec.jpg, Pink pupa due to pupation among inanimate objects

Adult

Male and female ''D. chrysippus'' butterflies look very similar and are also similar in size. Adult butterflies typically have a wingspan of . The bodies of adult plain tigers are about long, and their antennae are about long. Depending on temperature, males live about 10–15 days and females live about 7–12 days. File:Newly emerged Plain tiger female.JPG, A newly emerged plain tiger female in captivity File:Plain tiger m up.JPG, Male ''D. c. chrysippus'', recently emerged from chrysalis File:Plain tiger form Alcipoides from Karanji lake Mysore IMG 7834.jpg, Form ''alcippoides'' in

Mysore Mysore (), officially Mysuru (), is a city in the southern part of the state of Karnataka, India. Mysore city is geographically located between 12° 18′ 26″ north latitude and 76° 38′ 59″ east longitude. It is located at an altitude of ...

Karnataka Karnataka (; ISO: , , also known as Karunāḍu) is a state in the southwestern region of India. It was formed on 1 November 1956, with the passage of the States Reorganisation Act. Originally known as Mysore State , it was renamed ''Karnat ...

, India File:Plain Tiger (Danaus chrysippus)- mud-puddling in Hyderabad, AP W IMG 9741.jpg, Mud-puddling File:Plain Tiger (Danaus chrysippus), Bangalore, India.jpg, Resting with its wings closed File:Danaus chrysippus02.jpg, Basking

Enemies

Predators

Egg and larval predators

Most predators of the early developmental stages of ''D. chrysippus'' are

arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...

s. Such potential predators include various kinds of

spider Spiders ( order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species ...

assassin bugs 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 exam ...

cockroach Cockroaches (or roaches) are a Paraphyly, paraphyletic group of insects belonging to Blattodea, containing all members of the group except termites. About 30 cockroach species out of 4,600 are associated with human habitats. Some species are we ...

es,

ladybugs Coccinellidae () is a widespread family of small beetles ranging in size from . They are commonly known as ladybugs in North America and ladybirds in Great Britain. Some entomologists prefer the names ladybird beetles or lady beetles as they ...

ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of ...

s, and

mantises 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 hav ...

. The caterpillars will even cannibalize each other. Egg and larval mortality is often high; as many as 84% of eggs may be lost to predation and up to 97% of larvae can be lost by the fifth instar, although most larval deaths occur during the third instar.

Adult predators

The most common predator of adult plain tigers are

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 lightweig ...

s. In eastern Africa, the most common predator is the fiscal shrike ''L. c. humeralis''.

Parasites

There are several

organism In biology, an organism () is any living system that functions as an individual entity. All organisms are composed of cells (cell theory). Organisms are classified by taxonomy into groups such as multicellular animals, plants, and ...

s which parasitize the larvae of ''D. chrysippus''. The fly ''S. flavohalterata'' of the

Family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...

Tachinidae The Tachinidae are a large and variable family of true fly, flies within the insect order Fly, Diptera, with more than 8,200 known species and many more to be discovered. Over 1,300 species have been described in North America alone. Insects in t ...

'' is responsible for small amounts of parasitization in ''D. chrysippus'' populations. It is unclear whether the fly oviposits on the eggs of ''D. chrysippus'' or whether the fly oviposits on leaves which are then consumed by ''D. chrysippus'' larvae. ''S. flavohalterata'' does not kill the larvae, and development is normal until the pupal stage, when larvae dies and the parasite emerges from the pupa instead. ''A. chrysippi'', a parasitic

wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. ...

of the

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 ...

, oviposits on larvae early in their development and then kills them in the later stages. As many as fifty wasps may emerge from one large caterpillar, and they then pupate on the deceased host. Parasitic wasps of the genus ''Charops'' also infest plain tiger

s, likely during the egg or first instar stage, and then kill the larvae in a later instar stage. ''Sturmia convergens'' is also a parasitoid of ''D. chrysippus.''

Diseases

The plain tiger is infected by a male-killing

bacterium Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were amon ...

called ''

Spiroplasma ''Spiroplasma'' is a genus of Mollicutes, a group of small bacteria without cell walls. ''Spiroplasma'' shares the simple metabolism, parasitic lifestyle, fried-egg colony morphology and small genome of other ''Mollicutes'', but has a distinctiv ...

''. Male-killing bacteria are transmitted vertically, from mother to offspring. Female plain tigers infected with ''Spiroplasma'' will produce all-female broods, because the bacteria kills infected male offspring during either their embryonic or first larval

stage. Although male-killing bacteria are uncommon in species which lay eggs singly, experimentally treating infected females with

antibiotics An antibiotic is a type of antimicrobial substance active against bacteria. It is the most important type of antibacterial agent for fighting bacterial infections, and antibiotic medications are widely used in the treatment and prevention o ...

restored an even

sex ratio The sex ratio (or gender ratio) is usually defined as the ratio of males to females in a population. As explained by Fisher's principle, for evolutionary reasons this is typically about 1:1 in species which reproduce sexually. Many species d ...

to their subsequent broods, thus indicating that it is indeed ''Spiroplasma'' which is responsible for all-female broods in ''D. chrysippus''. However, the

prevalence In epidemiology, prevalence is the proportion of a particular population found to be affected by a medical condition (typically a disease or a risk factor such as smoking or seatbelt use) at a specific time. It is derived by comparing the number o ...

of this bacteria in the plain tiger seems to be restricted to east African populations.

Protective coloration and behavior

The plain tiger is mimicked by several species due to its unpalatability to potential predators. Previously, it was thought that

s obtained from food sources during the larval stage were responsible for the aversive nature of adult ''D. chrysippus'', but many larval food sources lack cardenolides, and some adult West African populations of ''D. chrysippus'' do not store cardenolides well, yet still repel predators. More recently, pyrrolizidine alkaloids have been proposed to be also responsible for the unpalatability of ''D. chrysippus''. Adult male danaines often feed on plants containing pyrrolizidine alkaloids, and although females rarely do, they may be protected simply through their resemblance to males of the same species. The ability of ''D. chrysippus'' to store cardenolides varies across populations, so likely both cardenolides and pyrrolizidine alkaloids contribute to the unpalatability of ''D. chrysippus'' to different extents depending on the population. Because the plain tiger is

(also called inedible), they are

aposematic Aposematism is the advertising by an animal to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defences which make the prey difficult to kill and eat, such as toxicity, venom, foul taste o ...

- their bright coloration serves as a warning to predators that they are either distasteful or

toxic Toxicity is the degree to which a chemical substance or a particular mixture of substances can damage an organism. Toxicity can refer to the effect on a whole organism, such as an animal, bacterium, or plant, as well as the effect on a subs ...

. Consequently, once a predator has made the mistake of attempting to eat a plain tiger, they will refrain in the future from attacking similarly colored butterflies. This has led to the

evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...

of a number of other species which mimic the plain tiger in order to co-opt the protection conferred by such bright coloration.

Batesian mimicry

Batesian mimics are

palatable Palatability (or palatableness) is the hedonic reward (i.e., pleasure) provided by foods or fluids that are agreeable to the "palate", which often varies relative to the homeostatic satisfaction of nutritional, water, or energy needs. The palatabi ...

species which mimic unpalatable species, and ''D. chrysippus'' is a model for several Batesian mimics, including '' H. misippus, P. poggei, M. marshalli'', and '' P. dardanus'' in east Africa. Batesian mimicry is only effective so long as the mimic is less common than the model, or predators will learn that the mimics are in fact edible and then attempt to eat the similar-looking unpalatable butterflies.

Müllerian mimicry

Müllerian mimicry Müllerian mimicry is a natural phenomenon in which two or more well-defended species, often foul-tasting and sharing common predators, have come to mimic each other's honest warning signals, to their mutual benefit. The benefit to Mülleria ...

occurs when multiple species which are all unpalatable evolve to resemble one another. In this case, the relative abundance of each species does not have a deleterious effect on any others, because a predator which eats any one of them will be deterred from eating any similar-looking butterflies. In

Uganda }), is a landlocked country in East Africa. The country is bordered to the east by Kenya, to the north by South Sudan, to the west by the Democratic Republic of the Congo, to the south-west by Rwanda, and to the south by Tanzania. The sou ...

, ''D. chrysippus'' has several Müllerian mimics, including '' A. encedon'' and '' A. encedana''.

Genetics

Subspecies

Despite the external similarity, the common tiger (''D. genutia'') is not closely related to the plain tiger. Three subspecies were considered valid in a 2005 review:Smith ''et al.'' (2005) * ''Danaus chrysippus chrysippus'' : Asia,

Mediterranean The Mediterranean Sea is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean Basin and almost completely enclosed by land: on the north by Western Europe, Western and Southern Europe and Anatolia, on the south by North Africa ...

region, northern tropical Africa * ''Danaus chrysippus alcippus'' (Cramer, 1777) - formerly ''D. c. aegyptius'' : From the

Cape Verde Islands , national_anthem = () , official_languages = Portuguese , national_languages = Cape Verdean Creole , capital = Praia , coordinates = , largest_city = capital , demonym ...

through tropical Africa to

Yemen Yemen (; ar, ٱلْيَمَن, al-Yaman), officially the Republic of Yemen,, ) is a country in Western Asia. It is situated on the southern end of the Arabian Peninsula, and borders Saudi Arabia to the Saudi Arabia–Yemen border, north and ...

and

Oman Oman ( ; ar, عُمَان ' ), officially the Sultanate of Oman ( ar, سلْطنةُ عُمان ), is an Arabian country located in southwestern Asia. It is situated on the southeastern coast of the Arabian Peninsula, and spans the mouth of ...

. Browner with broader white forewing spots. * ''Danaus chrysippus orientis'' (Aurivillius, 1909) - formerly ''D. c. liboria'' : Saint Helena, southern tropical Africa to

South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. It is bounded to the south by of coastline that stretch along the Atlantic Ocean, South Atlantic and Indian Oceans; to the ...

, Comoros,

Seychelles Seychelles (, ; ), officially the Republic of Seychelles (french: link=no, République des Seychelles; Creole: ''La Repiblik Sesel''), is an archipelagic state consisting of 115 islands in the Indian Ocean. Its capital and largest city, ...

and

Mascarenes The Mascarene Islands (, ) or Mascarenes or Mascarenhas Archipelago is a group of islands in the Indian Ocean east of Madagascar consisting of the islands belonging to the Republic of Mauritius as well as the French department of Réunion. Thei ...

. Small white forewing spots. ''D. c. alcippus'' is well on the way of becoming a distinct species. On the other hand, the former subspecies ''petilia'' is nowadays recognized as a good species, the lesser wanderer (''D. petilia''). More enigmatic is the status of the former subspecies (or forms) ''dorippus'' and ''bataviana''. These are tentatively also regarded as a distinct species, the dorippus tiger (''D. dorippus''). However, it appears (from analysis of

mtDNA Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial DNA ...

sequence In mathematics, a sequence is an enumerated collection of objects in which repetitions are allowed and order matters. Like a set, it contains members (also called ''elements'', or ''terms''). The number of elements (possibly infinite) is calle ...

s, which are only inherited from the mother) that the dorippus tiger is the product of an ancient lineage of ''Danaus'' hybridizing with plain tiger females. As the plain tiger is known to be parasitized at least occasionally by ''

bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of prokaryotic microorganisms. Typically a few micrometr ...

which selectively kill off male hosts,Jiggins ''et al.'' (2000) a subsequent scarcity of plain tiger males might have led to this hybridization and the

of the dorippus tiger. From the colour pattern of this species, it can be assumed that the ancient lineage had no black apex on the forewings, as this characteristic is still absent in ''D. dorippus''. The presumed subspecies ''cratippus'' most likely belongs to either the lesser wanderer or the dorippus tiger, but confirmation of its taxonomic status requires more research. In any case, these three species are closely related; their closest relatives, in turn, might be the

soldier A soldier is a person who is a member of an army. A soldier can be a conscripted or volunteer enlisted person, a non-commissioned officer, or an officer. Etymology The word ''soldier'' derives from the Middle English word , from Old French ...

(''D. eresimus'') and

queen Queen or QUEEN may refer to: Monarchy * Queen regnant, a female monarch of a Kingdom ** List of queens regnant * Queen consort, the wife of a reigning king * Queen dowager, the widow of a king * Queen mother, a queen dowager who is the mother ...

(''D. gilippus'') butterflies. Several local forms have been described from Asia: * ''Danaus chrysippus chrysippus'' f. ''alcippoides'' : The upper hindwing is more or less very white; about half of the individuals have a second submarginal spot in the forewing. Occasionally found in South-East Asia, very rarely in India. * ''Danaus chrysippus chrysippus'' f. ''gelderi'' : The upper hindwing has white markings. Occasionally found on Sulawesi. * ''Danaus chrysippus chrysippus'' f. ''bowringi'' : The upper hindwing has a subapical band composed of somewhat larger spots, and an additional forewing spot as in f. ''alcippoides'' is always present. Found throughout the eastern parts of this subspecies' range. On the other hand, the plethora of named

taxa In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...

from Africa are apparently F₁ or F₂ hybrids between the plain tiger subspecies (the contact zone of which is in the general area of

) and/or ''D. dorippus'': * ''Danaus chrysippus'' × ''alcippoides'' : is ''D. c. chrysippus'' × ''D. c. alcippus'' * ''Danaus'' × ''transiens'', ''Danaus'' × ''klugii'', ''Danaus'' × ''albinus'' and ''Danaus'' × ''semialbinus'' : are ''D. c. alcippus'' × ''D. dorippus''

Genomes

When ''D. chrysippus'' was analyzed via a sample from Kampala, Uganda, it was found that the population was undergoing a significant level of evolutionary change. Three loci were examined, and genotypic frequency differences found at two of the three suggested that opposing selective forces, likely pertaining to Mullerian and Batesian mimicry, acting on males and females is contributing to a balanced polymorphism.

Courtship and mating

Plain Tiger (Danaus chrysippus) mating in Hyderabad, AP W IMG 9493

In addition to conferring protection from predators, pyrrolizidine alkaloids are necessary in the mating ritual of ''D. chrysippus''. Male plain tigers use the alkaloids to synthesize

pheromone 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 ...

s which are stored in

hair-pencil Hair-pencils and coremata are pheromone signaling structures present in lepidopteran males. Males use hair-pencils in courtship behaviors with females. The pheromones they excrete serve as both aphrodisiacs and tranquilizers to females as well as r ...

s sheathed in alar organs, which are specialized scales on top of the hindwing. The hair-pencils are fanned out during courtship to release these pheromones, and this appears to be necessary for attracting females. Males deprived of pyrrolizidine alkaloids in their diet are considerably less successful in mating; mating appears to occur preferentially between butterflies of the same subspecies, so coloration is likely also an important signal in the mating process. Female plain tigers have been recorded as mating up to four times.

References

* * * * * Gil-T., Felipe (2006): A new hostplant for ''Danaus plexippus'' L. in Europe. A study of cryptic preimaginal polymorphism within ''Danaus chrysippus'' L. in southern Spain (Andalusia) (Lepidoptera, Nymphalidae, Danainae). ISSN 0171-0079 , ''Atalanta'' 37 (1/2): 143–149, 279. Full article

* HOSTS (2007)
A Database of the World's Lepidopteran Hostplants
Retrieved 2007-AUG-01. * * * Larsen, Torben (1994)

'' Saudi Aramco World'', issue 5 (September/October): 24–27. * * *

External links

AdaMerOs - Butterflies Watching & Photography Society in Turkey

Sri Lanka Wild Information Database
{{Taxonbar, from=Q2663244 Danaus (butterfly) Butterflies of Africa Butterflies of Asia Butterflies of Europe Butterflies of Indochina Butterflies of Java Butterflies of Singapore Insects of Cape Verde Insects of Egypt Insects of Pakistan Art of ancient Egypt Butterflies described in 1758 Taxa named by Carl Linnaeus