''Hypolimnas bolina'', the great eggfly, common eggfly, varied eggfly, or in New Zealand the blue moon butterfly, is a species of

nymphalid 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 ...

butterfly Butterflies are winged insects from the lepidopteran superfamily Papilionoidea, characterized by large, often brightly coloured wings that often fold together when at rest, and a conspicuous, fluttering flight. The oldest butterfly fossi ...

found from

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 ...

Asia Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...

and

Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...

Appearance

Race ''bolina''

''H. bolina'' is a black-bodied butterfly with a wingspan of about . The species has a high degree of

sexual dimorphism Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...

. The female is

mimetic Mimesis (; , ''mīmēsis'') is a term used in literary criticism and philosophy that carries a wide range of meanings, including ''imitatio'', imitation, Similarity (philosophy), similarity, receptivity, representation (arts), representation, m ...

with multiple morphs.

Male

Males are monomorphic. The

dorsal Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage The fus ...

wing surface is jet black but features three prominent spots, two on the

forewing Insect wings are adult outgrowths of the insect exoskeleton that enable insects to fly. They are found on the second and third thoracic segments (the mesothorax and metathorax), and the two pairs are often referred to as the forewings and hindwi ...

and one on the

hindwing Insect wings are adult outgrowths of the insect exoskeleton that enable insects to fly. They are found on the second and third thoracic segments (the mesothorax and metathorax), and the two pairs are often referred to as the forewings and hindw ...

. To a human observer these appear as white spots fringed with blue-violet. They actually consist of a white center overlain by bright ultraviolet

iridescence Iridescence (also known as goniochromism) is the phenomenon of certain surfaces that appear gradually to change colour as the angle of view or the angle of illumination changes. Iridescence is caused by wave interference of light in microstru ...

, a colour generated by nanostructures on the wing scale surface. Numerous smaller white spots fringe the fore- and hindwings. The

ventral Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...

surface lacks any

ultraviolet Ultraviolet radiation, also known as simply UV, is electromagnetic radiation of wavelengths of 10–400 nanometers, shorter than that of visible light, but longer than X-rays. UV radiation is present in sunlight and constitutes about 10% of ...

iridescence and consists essentially of banded white markings set against a brownish background. These males exhibit lower body mass per unit wing area, implying lower wing loading, more elongate wings, resulting in a higher aspect ratio, which are also the characteristics used to differ them by their

conspecific 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 ...

females. They bear striking resemblance to ''

Hypolimnas alimena ''Hypolimnas alimena'', the blue-banded eggfly, is a species of butterfly in the family Nymphalidae. It is found in the Solomon Islands, Indonesia (Aru Islands Regency, Aru Islands, Irian Jaya Barat, Irian Jaya, Kai Islands, Kei Islands, Maluku I ...

'' in terms of

flight Flight or flying is the motion (physics), motion of an Physical object, object through an atmosphere, or through the vacuum of Outer space, space, without contacting any planetary surface. This can be achieved by generating aerodynamic lift ass ...

musculature Skeletal muscle (commonly referred to as muscle) is one of the three types of vertebrate muscle tissue, the others being cardiac muscle and smooth muscle. They are part of the somatic nervous system, voluntary muscular system and typically are a ...

(

thoracic The thorax (: thoraces or thoraxes) or chest is a part of the anatomy of mammals and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main ...

mass).

Female

Females are hugely variable due to the presence of both genetic polymorphism and phenotypic plasticity. Polymorphism is expressed primarily on the dorsal surface, with morphs varying in the presence of white, orange, and blue markings. One genetic morph, named '' euploeoides'' by Clarke & Sheppard (1975), is thought to present a mimic of one or several members of the genus ''

Euploea ''Euploea'' is a genus of milkweed butterflies. The species are generally dark in coloration, often quite blackish, for which reason they are commonly called crows. As usual for their subfamily, they are poisonous due to feeding on milkweeds a ...

''. The female ventral wing surfaces are similar to those of the male. Phenotypic plasticity is such that individuals are generally darker if they develop under cooler temperatures.

Gallery

Blue-moon butterfly (Hypolimnas bolina nerina) female underside Rarotonga.jpg, female ''H. p. nerina''
Cook Islands File:Common eggfly (Hypolimnas bolina naresii) female Viti Levu.jpg, female ''H. p. naresii''
Fiji Hypolimnas bolina by Kadavoor.jpg, Male, underside Eggs of Hypolimnas bolina Linnaeus, 1758 – Great Eggfly OP2A9916.jpg, Eggs Great eggfly cat.jpg, Caterpillar of race ''bolina'' Great eggfly pupa.jpg, Pupa of race ''bolina''

Distribution

''H. bolina'' is found in

in the west, through to

South South is one of the cardinal directions or compass points. The direction is the opposite of north and is perpendicular to both west and east. Etymology The word ''south'' comes from Old English ''sūþ'', from earlier Proto-Germanic ''*sunþa ...

and

Southeast Asia Southeast Asia is the geographical United Nations geoscheme for Asia#South-eastern Asia, southeastern region of Asia, consisting of the regions that are situated south of China, east of the Indian subcontinent, and northwest of the Mainland Au ...

Cambodia Cambodia, officially the Kingdom of Cambodia, is a country in Southeast Asia on the Mainland Southeast Asia, Indochinese Peninsula. It is bordered by Thailand to the northwest, Laos to the north, and Vietnam to the east, and has a coastline ...

, the South Pacific islands (

French Polynesia French Polynesia ( ; ; ) is an overseas collectivity of France and its sole #Governance, overseas country. It comprises 121 geographically dispersed islands and atolls stretching over more than in the Pacific Ocean, South Pacific Ocean. The t ...

Tonga Tonga, officially the Kingdom of Tonga, is an island country in Polynesia, part of Oceania. The country has 171 islands, of which 45 are inhabited. Its total surface area is about , scattered over in the southern Pacific Ocean. accordin ...

Tuvalu Tuvalu ( ) is an island country in the Polynesian subregion of Oceania in the Pacific Ocean, about midway between Hawaii and Australia. It lies east-northeast of the Santa Cruz Islands (which belong to the Solomon Islands), northeast of Van ...

Samoa Samoa, officially the Independent State of Samoa and known until 1997 as Western Samoa, is an island country in Polynesia, part of Oceania, in the South Pacific Ocean. It consists of two main islands (Savai'i and Upolu), two smaller, inhabited ...

Vanuatu Vanuatu ( or ; ), officially the Republic of Vanuatu (; ), is an island country in Melanesia located in the South Pacific Ocean. The archipelago, which is of volcanic origin, is east of northern Australia, northeast of New Caledonia, east o ...

, and

Fiji Fiji, officially the Republic of Fiji, is an island country in Melanesia, part of Oceania in the South Pacific Ocean. It lies about north-northeast of New Zealand. Fiji consists of an archipelago of more than 330 islands—of which about ...

), and occurs in parts of

as far south as Victoria during summer and autumn,

Japan Japan is an island country in East Asia. Located in the Pacific Ocean off the northeast coast of the Asia, Asian mainland, it is bordered on the west by the Sea of Japan and extends from the Sea of Okhotsk in the north to the East China Sea ...

Taiwan Taiwan, officially the Republic of China (ROC), is a country in East Asia. The main geography of Taiwan, island of Taiwan, also known as ''Formosa'', lies between the East China Sea, East and South China Seas in the northwestern Pacific Ocea ...

, and

New Zealand New Zealand () is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and List of islands of New Zealand, over 600 smaller islands. It is the List of isla ...

. Appearances in New Zealand appear to be linked with favourable winds during April–June migratory periods in Australia, with the butterfly being recorded in 1956, 1971, and 1995. However, there are

iNaturalist iNaturalist is an American 501(c)(3) nonprofit social network of naturalists, citizen scientists, and biologists built on the concept of mapping and sharing observations of biodiversity across the globe. iNaturalist may be accessed via its web ...

records for

for every year since 2013. No populations have established locally in

Habitat

''H. bolina'' is a generally common butterfly across most tropical and sub-tropical habitat types, including wet/dry woodland (such as

tropical savanna Tropical and subtropical grasslands, savannas, and shrublands is a terrestrial biome defined by the World Wide Fund for Nature. The biome is dominated by grass and/or shrubs located in semi-arid to semi-humid climate regions of subtropical and t ...

rainforest Rainforests are forests characterized by a closed and continuous tree Canopy (biology), canopy, moisture-dependent vegetation, the presence of epiphytes and lianas and the absence of wildfire. Rainforests can be generally classified as tropi ...

, and

shrubland Shrubland, scrubland, scrub, brush, or bush is a plant community characterized by vegetation dominance (ecology), dominated by shrubs, often also including grasses, herbaceous plant, herbs, and geophytes. Shrubland may either occur naturally o ...

. It is a common visitor to suburban back yards and other areas of human disturbance. In the Australian tropics, ''H. bolina'' is particularly common along tracks, streambeds, and the corridors formed between remnant

gallery forest A gallery forest is one formed as a corridor along rivers or wetlands, projecting into landscapes that are otherwise only sparsely treed such as savannas, grasslands, or deserts. The gallery forest maintains a more temperate microclimate above th ...

and cultivated sugar cane fields. Both sexes are invariably found in association with one or more larval hostplant species, especially disturbance species such as '' Synedrella nodiflora'', ''

Sida rhombifolia ''Sida rhombifolia'', commonly known as arrowleaf sida, is a perennial plant, perennial or sometimes annual plant, annual plant in the Family Malvaceae, native to the Old World tropics and subtropics. Other common names include rhombus-leaved sid ...

'', and '' Commelina cyanea''.

Mimicry

To the west the female is monomorphic, mimicking species of the oriental and Australasian danaid genus ''

''. Eastwards ''H. bolina'' is frequently polymorphic and most forms are then non-mimetic. In areas where it resembles ''Euploea'' the butterfly has usually been designated a Batesian mimic.

Life cycle and ecology

Males are notably territorial. Individuals are known to return to defend the same location for up to 54 days, with site fidelity increasing with age. Territories that enhance the visual detection of adult females are preferred. Males primarily utilize sit-and-wait strategy for locating potential mates. Females are usually found gliding close to the ground in southern areas. Unlike that of ''H. alimena,'' these females possess a strong preference towards exaggerated visual signals (dorsal blue coloration). Unlike congenerics such as '' Hypolimnas anomala'', female ''H. bolina'' most often lay one or two eggs at a time. These are typically laid on the leaf underside. Early season (post-diapause) females in the Australian wet-dry tropics target freshly germinated seedlings of their favoured host in this region, '' Synedrella nodiflora''. Eggs hatch after 3 days under a constant incubation temperature of ca. 25 degrees C. Newly hatched larva first consume their egg shell before feeding on the leaf upon which they were laid. Later larval instars are highly mobile and readily disperse in search of new host foliage. Individuals rarely pupate on (or in the immediate vicinity of) their host plant.

Host plants

Race ''bolina'' breeds on ''

'', '' Elatostemma cuneatum'', ''

Portulaca oleracea ''Portulaca oleracea'' (common purslane, also known as little hogweed, or pursley) is a succulent plant in the family Portulacaceae. Description The plant may reach in height. It has smooth, reddish, mostly prostrate stems, and the leaves, w ...

'', '' Laportea interrupta'', '' Triumfetta pentandra'', and '' Asystasia'' species. Other hosts include '' Elatostema cuneatum'', '' Fleurya interrupta'', '' Pseuderanthemum variabile'', ''

Ipomoea batatas The sweet potato or sweetpotato (''Ipomoea batatas'') is a dicotyledonous plant in the morning glory family, Convolvulaceae. Its sizeable, starchy, sweet-tasting tuberous roots are used as a root vegetable, which is a staple food in parts of the ...

,'' ''

Alternanthera denticulata ''Alternanthera denticulata'' (common name lesser joyweed) is a small prostrate white-flowering herb in the Amaranthaceae family. It is native to all states and territories of Australia, New Guinea, and the North Island of New Zealand. It is al ...

'', and '' Synedrella nodiflora''. Caterpillars accept leaves of Pipiturus argenteus. They are also known to feed on ''

Urtica dioica ''Urtica dioica'', often known as common nettle, burn nettle, stinging nettle (although not all plants of this species sting) or nettle leaf, or just a nettle or stinger, is a herbaceous perennial flowering plant in the family Urticaceae. Or ...

'' and ''

Malva ''Malva'' is a genus of herbaceous annual, biennial, and perennial plants in the family Malvaceae. It is one of several closely related genera in the family to bear the common English name mallow. The genus is widespread throughout the temp ...

'' species. While in New Zealand, adult butterflies have been observed feeding on the nectar of ''

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

'', ''

Grevillea ''Grevillea'' (), commonly known as spider flowers, is a genus of about 360 species of evergreen flowering plants in the family Proteaceae. Plants in the genus ''Grevillea'' are shrubs, rarely trees, with the leaves arranged alternately along the ...

'', '' Hebe'', ''

Symphyotrichum novi-belgii ''Symphyotrichum novi-belgii'' (formerly ''Aster novi-belgii''), commonly called New York aster, is a species of flowering plant. It is the type species for ''Symphyotrichum'', a genus in the family Asteraceae, whose species were once consider ...

'', and ''

Solanum muricatum ''Solanum muricatum'' is a species of evergreen shrub native to South America and grown for its sweet edible fruit. It is known as ''pepino dulce'' ("sweet cucumber" in English, in order to differentiate it from cucumber which is also called " ...

'' plants.

Eggs

The eggs are a pale, glassy green with longitudinal ridges except on the top.

Caterpillar

After about four days the eggs hatch. The

caterpillar 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 ...

s immediately disperse. They are black with an orange head. The last segment is also orange. The head bears a pair of long branched black horns. The body surface is also covered with long, branched, orangish black spines. These spines look whitish and transparent immediately after

moulting In biology, moulting (British English), or molting (American English), also known as sloughing, shedding, or in many invertebrates, ecdysis, is a process by which an animal casts off parts of its body to serve some beneficial purpose, either at ...

, but soon become the usual orange. In later

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 ...

s the spiracles are surrounded by thin, dirty orange rings. Infection by ''

Wolbachia ''Wolbachia'' is a genus of gram-negative bacteria infecting many species of arthropods and filarial nematodes. The symbiotic relationship ranges from parasitism to obligate mutualism. It is one of the most common parasitic microbes of arthrop ...

'' bacteria is known to exclusively kill male specimens.

Pupa

The pupa is suspended by just one point. It is brown with a grey tinge on the wings. The abdominal segments have distinct tubercles. The surface of the pupa is rough. The butterfly emerges after seven to eight days as pupae (female development is always a bit longer).

Recent evolutionary changes

On the

Samoan Islands The Samoan Islands () are an archipelago covering in the central Pacific Ocean, South Pacific, forming part of Polynesia and of the wider region of Oceania. Political geography, Administratively, the archipelago comprises all of the Samoa, Indep ...

Upolu Upolu is an island in Samoa, formed by a massive basaltic shield volcano which rises from the seafloor of the western Pacific Ocean. The island is long and in area, making it the second largest of the Samoan Islands by area. With approximate ...

and Savai'i, a ''

'' strain wBol1 had been killing the male members of ''Hypolimnas bolina''. The problem was so severe that by 2001, males made up only 1% of the population. However, in 2007, it was reported that within a span of just 10 generations (about 5 years), the males had evolved to develop immunity to the parasite, and the male population increased to nearly 40%. This evolutionary event involved changes at a single genomic region on chromosome 25, and represents one of the fastest examples of natural selection observed to date in natural populations. Ed Yong has written a popular science account of this highly unusual evolutionary event. ''Wolbachia'' derived from wBol1 have recently been observed in ''H. bolina'' from Java, Indonesia, to be insensitive to suppression, carrying an additional prophage element with additional male-killing determinants. This observation indicated an escalating arms-race may be operating in this species

Subspecies

Listed alphabetically:"''Hypolimnas'' Hübner, [1819]"
at Markku Savela's ''Lepidoptera and Some Other Life Forms''. *''H. b. bolina'' (Linnaeus, 1758) – (Sumatra, Java, Lesser Sunda Islands, western Borneo, Sulawesi, Salayar, Kabaena, Galla, Banggai, Sula, Maluku, New Guinea, Solomon Islands, Australia, New Caledonia) *''H. b. constans'' (Butler, 1875) – (Tasmania?) *''H. b. enganica'' Fruhstorfer, 1904 – (Engano Island) *''H. b. gigas'' (Oberthür, 1879) – (Sangihe) *''H. b. incommoda'' Butler, 1879 *''H. b. inconstans'' Butler, 1873 – (Navigator Islands) *''H. b. jacintha'' (Drury, 1773) *''H. b. jaluita'' Fruhstorfer, 1903 *''H. b. kezia'' (Butler) – (Formosa) *''H. b. kraimoku'' (Eschscholtz, 1821) – (Lifu) *''H. b. labuana'' Butler, 1879 – (Labuan) *''H. b. lisianassa'' (Cramer, 1779) – (Moluccas) *''H. b. listeri'' Butler, 1888 – (Christmas Island) *''H. b. montrouzieri'' (Butler) – (Woodlark, Fergusson, Trobriand Islands) *''H. b. naresii'' Butler, 1883 – (Fiji) *''H. b. nerina'' (Fabricius, 1775) – (Timor - Kai, Aru, Waigeu, West Irian - Papua, northern Australia - eastern Victoria, Bismarck Archipelago, Solomon Islands, New Zealand) *''H. b. pallescens'' (Butler) – (Fiji) *''H. b. philippensis'' (Butler, 1874) – (Philippines) *''H. b. pulchra'' (Butler) – (New Caledonia) *''H. b. rarik'' Eschscholtz, 1821) – (Lifu)

References

External links

Evolution of Male-Killer Suppression in a Natural Population

(English/German)
''Hypolimnas bolina''
at CalPhotos {{Taxonbar">from=Q309627 bolina Butterflies of Indochina Butterflies of Taiwan Butterflies of Singapore">Butterflies of Taiwan">Butterflies of Indochina Butterflies of Taiwan Butterflies of Singapore Butterflies of New Zealand Butterflies described in 1758 Taxa named by Carl Linnaeus Articles containing video clips Fauna of Niue Butterflies of Japan Butterflies of Africa