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''Kallima inachus'', the orange oakleaf, Indian oakleaf or dead leaf, is a
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 a red ...
butterfly found in
Tropical Asia Tropical Asia refers to the entirety of the areas in Asia with a tropical climate. These areas are of geographic and economic importance due to their natural resources and biodiversity, which include many species of agricultural value. There are ...
from
India India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the seventh-largest country by area, the second-most populous country, and the most populous democracy in the world. Bounded by the Indian Ocean on the so ...
to
Japan Japan ( ja, 日本, or , and formally , ''Nihonkoku'') is an island country in East Asia. It is situated in the northwest Pacific Ocean, and is bordered on the west by the Sea of Japan, while extending from the Sea of Okhotsk in the north ...
. With wings closed, it closely resembles a dry leaf with dark veins and is a commonly cited example of
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 ...
.


Description

The butterfly wings are shaped like a leaf when in the closed position. When the wings are closed, only the
cryptic Cryptic may refer to: In science: * Cryptic species complex, a group of species that are very difficult to distinguish from one another * Crypsis, the ability of animals to blend in to avoid observation * Cryptic era, earliest period of the Earth ...
underside markings are visible, which consists of irregular patterns and striations in many shades of biscuit, buff, browns, yellow, and black. The veins are darkened and resemble the veins of a leaf. The resemblance to a dried leaf, a
masquerade Masquerade or Masquerader may refer to: Events * Masquerade ball, a costumed dance event * Masquerade ceremony, a rite or cultural event in many parts of the world, especially the Caribbean and Africa * Masqueraders, the performers in the West ...
, is extremely realistic and gives the genus its common names, the oakleaf or dead leaf. When the wings are open, the forewing exhibits a black apex, an orange discal band and a deep blue base. There are two white
oculi An oculus (; ) is a circular opening in the center of a dome or in a wall. Originating in antiquity, it is a feature of Byzantine and Neoclassical architecture. It is also known as an '' œil-de-boeuf'' from the French, or simply a "bull's-e ...
, one along the margin of the apical black band, and the other bordering the orange and deep blue areas. The hindwing is more uniformly blue but diffused with brown patches along the
termen Termen is a municipality in the district of Brig in the canton of Valais in Switzerland. History Termen is first mentioned in 1201 as ''Terman''. Geography Termen has an area, , of . Of this area, 42.2% is used for agricultural purposes, whil ...
. Male and female butterflies are similar except that the female is generally larger and has the
apex The apex is the highest point of something. The word may also refer to: Arts and media Fictional entities * Apex (comics), a teenaged super villainess in the Marvel Universe * Ape-X, a super-intelligent ape in the Squadron Supreme universe *Apex ...
of 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 hindwin ...
protrude to form a longer point. Females also tend to be more reddish on the underside and the yellow mottled markings tend to be paler. The butterfly exhibits
polyphenism A polyphenic trait is a trait for which multiple, discrete phenotypes can arise from a single genotype as a result of differing environmental conditions. It is therefore a special case of phenotypic plasticity. There are several types of polyphen ...
, i.e. there are specific dry-season and wet-season forms which differ in colouration and size; the wet-season form tends to be smaller. The wingspan of the butterfly ranges from . Detailed description as given in Bingham (1905).


Dry-season form

The forewing discoidal cell, interspace 1a, 1 to near apex, basal half of 2, and extreme bases of 3 and 4 rich violet blue, the borders of the discocellulars and the interspaces of veins 2, 3 and 4 are black, spread diffusely outwards in interspaces 1a and 1. A very broad oblique discal orange band from costa to apices of interspaces 1 and 2, this orange band is sprinkled with bluish black scales; apical third of wing velvety purpurescent (purple) black; a hyaline (glass-like) transverse spot near middle of interspace 2, and a subtriangular similar small preapical spot. Hindwing more uniform violescent blue; the costal margin and apex very broadly brown, somewhat densely irrorated (sprinkled) with dusky violescent black scales; dorsal margin brown; a ridge of long brownish hairs along vein 1 spreading on to the dorsal margin. Forewings and hindwings crossed by a subterminal dusky zigzag line commencing about the middle of interspace 3 in the forewing, and most conspicuous on the hindwing. Underside very closely resembles a dry leaf; ground colour very variable, but usually some shade of brown (rusty, greyish, and yellowish browns being the most common), always with scattered dark dots or little dark patches having the appearance of fungus-like or lichenous growths so common on dead leaves in the tropics. When the insect closes its wings over its back the likeness to a dead leaf is most striking, and is heightened by a straight transverse, narrow, dark band running from the apex of the forewing to the tornus of the hindwing, often with oblique narrower similar bands or lines given off from it, all simulating very closely the midrib and lateral veins of a leaf. The hindwing in all specimens has a more or less obsolescent or faint series of postdiscal
ocelli A simple eye (sometimes called a pigment pit) refers to a form of eye or an optical arrangement composed of a single lens and without an elaborate retina such as occurs in most vertebrates. In this sense "simple eye" is distinct from a multi-l ...
, traces of which are also apparent on the forewing. Antennae dark brown; head, thorax, and abdomen dark violescent brown; beneath, the palpi, thorax and abdomen paler earthy brown.


Wet-season form

Smaller than the dry-season form, but very similar to it. The colours are richer and darker, and the orange discal band more broadly bordered with black on the inner side. On the underside some of the specimens from areas of heavy rainfall have the ground colour very dark ochraceous brown. File:Kallima inachus OPENWING.jpg File:Orange Oakleaf (Kallima inachus) at Samsing, Duars, West Bengal W IMG 6241.jpg, Dry-season, female, upperside File:Kallima_inachis_formosana.jpg, Dry-season, female, underside File:Orange oak leaf in Pakke Tiger Reserve.JPG, Wet-season, female, upperside File:Orange_Oakleaf_(Kallima_inachus).jpg, Wet-season, female, underside File:Indian oakleaf or dead leaf DSCN2192 11.jpg, Dry-season at
Sattal Sattal or Sat Tal (Hindi for "seven lakes") is an interconnected group of seven freshwater lakes situated in the Lower Himalayan Range near Bhimtal, a town of the Nainital district in Uttarakhand, India. During the British Raj, the area had a t ...


Distribution

The orange oakleaf is found in
India India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the seventh-largest country by area, the second-most populous country, and the most populous democracy in the world. Bounded by the Indian Ocean on the so ...
,
Nepal Nepal (; ne, नेपाल ), formerly the Federal Democratic Republic of Nepal ( ne, सङ्घीय लोकतान्त्रिक गणतन्त्र नेपाल ), is a landlocked country in South Asia. It is mai ...
,
Bhutan Bhutan (; dz, འབྲུག་ཡུལ་, Druk Yul ), officially the Kingdom of Bhutan,), is a landlocked country in South Asia. It is situated in the Eastern Himalayas, between China in the north and India in the south. A mountainous ...
,
Bangladesh Bangladesh (}, ), officially the People's Republic of Bangladesh, is a country in South Asia. It is the eighth-most populous country in the world, with a population exceeding 165 million people in an area of . Bangladesh is among the mos ...
, and
Myanmar Myanmar, ; UK pronunciations: US pronunciations incl. . Note: Wikipedia's IPA conventions require indicating /r/ even in British English although only some British English speakers pronounce r at the end of syllables. As John C. Wells, Joh ...
, down to
Tenasserim Hills The Tenasserim Hills or Tenasserim Range ( my, တနင်္သာရီ တောင်တန်း, ; th, ทิวเขาตะนาวศรี, , ; ms, Banjaran Tanah Seri/Banjaran Tenang Sari) is the geographical name of a r ...
. In Southeast Asia it occurs in southern
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
,
Thailand Thailand ( ), historically known as Siam () and officially the Kingdom of Thailand, is a country in Southeast Asia, located at the centre of the Indochinese Peninsula, spanning , with a population of almost 70 million. The country is bo ...
,
Laos Laos (, ''Lāo'' )), officially the Lao People's Democratic Republic ( Lao: ສາທາລະນະລັດ ປະຊາທິປະໄຕ ປະຊາຊົນລາວ, French: République démocratique populaire lao), is a socialist ...
,
Taiwan Taiwan, officially the Republic of China (ROC), is a country in East Asia, at the junction of the East and South China Seas in the northwestern Pacific Ocean, with the People's Republic of China (PRC) to the northwest, Japan to the nort ...
, and
Vietnam Vietnam or Viet Nam ( vi, Việt Nam, ), officially the Socialist Republic of Vietnam,., group="n" is a country in Southeast Asia, at the eastern edge of mainland Southeast Asia, with an area of and population of 96 million, making i ...
. It has been also recorded from Pakistan in 2000. In India, the butterfly flies in the
Himalayas The Himalayas, or Himalaya (; ; ), is a mountain range in Asia, separating the plains of the Indian subcontinent from the Tibetan Plateau. The range has some of the planet's highest peaks, including the very highest, Mount Everest. Over 100 ...
at low elevations, from
Jammu and Kashmir Jammu and Kashmir may refer to: * Kashmir, the northernmost geographical region of the Indian subcontinent * Jammu and Kashmir (union territory), a region administered by India as a union territory * Jammu and Kashmir (state), a region administered ...
, through
Garhwal Garhwal may refer to the following topics associated with Uttarakhand, India: Places *Garhwal Himalaya, a sub-range of the Himalayas *Garhwal Kingdom, a former kingdom * Garhwal District (British Garhwal), a former district of British India *Gar ...
and
Kumaon Kumaon or Kumaun may refer to: * Kumaon division, a region in Uttarakhand, India * Kumaon Kingdom, a former country in Uttarakhand, India * Kumaon, Iran, a village in Isfahan Province, Iran * , a ship of the Royal Indian Navy during WWII See also ...
to
West Bengal West Bengal (, Bengali: ''Poshchim Bongo'', , abbr. WB) is a state in the eastern portion of India. It is situated along the Bay of Bengal, along with a population of over 91 million inhabitants within an area of . West Bengal is the fourt ...
,
Sikkim Sikkim (; ) is a state in Northeastern India. It borders the Tibet Autonomous Region of China in the north and northeast, Bhutan in the east, Province No. 1 of Nepal in the west and West Bengal in the south. Sikkim is also close to the Siligur ...
,
Arunachal Pradesh Arunachal Pradesh (, ) is a state in Northeastern India. It was formed from the erstwhile North-East Frontier Agency (NEFA) region, and became a state on 20 February 1987. It borders the states of Assam and Nagaland to the south. It shares int ...
, and other states of the northeast. It is also found in central and peninsular India; it flies in
Madhya Pradesh Madhya Pradesh (, ; meaning 'central province') is a state in central India. Its capital is Bhopal, and the largest city is Indore, with Jabalpur, Ujjain, Gwalior, Sagar, and Rewa being the other major cities. Madhya Pradesh is the seco ...
,
Jharkhand Jharkhand (; ; ) is a state in eastern India. The state shares its border with the states of West Bengal to the east, Chhattisgarh to the west, Uttar Pradesh to the northwest, Bihar to the north and Odisha to the south. It has an area of . It ...
,
Chhattisgarh Chhattisgarh (, ) is a landlocked state in Central India. It is the ninth largest state by area, and with a population of roughly 30 million, the seventeenth most populous. It borders seven states – Uttar Pradesh to the north, Madhya Prade ...
,
Orissa Odisha (English: , ), formerly Orissa ( the official name until 2011), is an Indian state located in Eastern India. It is the 8th largest state by area, and the 11th largest by population. The state has the third largest population of Sch ...
and
Andhra Pradesh Andhra Pradesh (, abbr. AP) is a state in the south-eastern coastal region of India. It is the seventh-largest state by area covering an area of and tenth-most populous state with 49,386,799 inhabitants. It is bordered by Telangana to the ...
; i.e. along the central Indian highlands to
Pachmarhi Pachmarhi is a hill station in Narmadapuram district of Madhya Pradesh states and territories of India, state of central India. It has been the location of a cantonment (Pachmarhi Cantonment) since British Raj. It is widely known as ''Satpura ...
and
Amarkantak Amarkantak (National Library at Kolkata romanization, NLK ''Amarakaṇṭaka'') is a pilgrim town and a Nagar Panchayat in Anuppur, Madhya Pradesh, India. The Amarkantak region is a unique natural heritage area and is the meeting point of the Vin ...
, the Western Ghats south to
Bhimashankar Bhimashankar Temple is a Jyotirlinga shrine located 50 km Khed taluka (alias Rajgurunagar), near Pune, in Maharashtra, India. It is located 127 km from Shivajinagar (in Pune) in the Ghat region of the Sahyadri Mountains. Bhīmāshan ...
, and in the
Eastern Ghats The Eastern Ghats are a discontinuous range of mountains along India's eastern coast. The Eastern Ghats pass through Odisha, Andhra Pradesh to Tamil Nadu in the south passing some parts of Karnataka as well as Telangana. They are eroded and cut ...
north of the river
Godavari The Godavari ( IAST: ''Godāvarī'' od̪aːʋəɾiː is India's second longest river after the Ganga river and drains into the third largest basin in India, covering about 10% of India's total geographical area. Its source is in Trimbakesh ...
. The status of the butterfly in India is "not rare", while in China, the butterfly is considered "rare". The orange oakleaf is encountered up to an altitude of in the hills; though
Mark Alexander Wynter-Blyth Mark Alexander Wynter-Blyth (15 August 1906 – 16 April 1963 Leysin, Switzerland) was an English schoolteacher and amateur naturalist who wrote one of the first field guides to the butterflies of the Indian region. He was also involved in censuses ...
records it as being encountered up to in regions of heavy rainfall in thickly forested mountainous and hilly regions. In the Kumaon Himalayas, ''K. inachus'' has been recorded to inhabit tropical deciduous forest between and subtropical evergreen forest above . In a survey of
Chongqing Chongqing ( or ; ; Sichuanese dialects, Sichuanese pronunciation: , Standard Mandarin pronunciation: ), Postal Romanization, alternately romanized as Chungking (), is a Direct-administered municipalities of China, municipality in Southwes ...
municipality,
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
carried out from 1998 to 2004, ''K. inachis'' was found to inhabit moist broad-leaf forests.


Habits

The orange oakleaf is a powerful flier and usually flies in dense forests with good rainfall, amongst undergrowth and along stream beds. It is attracted to tree sap and over-ripe fruit, and is also known to
mud-puddle Mud-puddling, or simply puddling, is a behaviour most conspicuous in butterflies, but occurs in other animals as well, mainly insects; they seek out nutrients in certain moist substances such as rotting plant matter, mud and carrion and they suck ...
. Much pursued by birds, when in danger the orange oakleaf flies erratically, soon dropping down into the foliage and occupying a stationary pose with wings closed, so that the birds are very often quite unable to find them. In such a pose, the butterfly resembles a dried leaf and is perfectly camouflaged. The natural enemies of the orange oakleaf include birds, ants, spiders, wasps (including ''
Trichogramma ''Trichogramma'' is a genus of minute polyphagous wasps that are endoparasitoids of insect eggs. ''Trichogramma'' is one of around 80 genera from the family Trichogrammatidae, with over 200 species worldwide.Consoli FL, Parra JRP, Zucchi RA (2010 ...
'' species), and some bacteria.


Life cycle

In the Himalayas, the butterfly is
multivoltine Voltinism is a term used in biology to indicate the number of broods or generations of an organism in a year. The term is most often applied to insects, and is particularly in use in sericulture, where silkworm varieties vary in their voltinism. ...
and flies from April to October. Kehimkar (2009) records the butterfly on the wing in India from April to December. In Chongqing one generation has been recorded as taking about 50 days from egg to imago. The egg period lasted about 6 days, the larval period 36 days, and involved 5 to 6
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 ass ...
s (usually 5) and with the pupation lasting about 10 days. The caterpillars bred successfully at temperatures of and relative humidity of 48 to 98%. The larval period could be reduced from 36 days in natural conditions to 16.8 to 23 days in captive breeding. In another study in China, in the Emei mountains (altitude ), the butterfly has three generations a year in which the first and second generations predominate. Most of the second generation, along with a few of the third and sometimes the first generation, go through the winter as
diapause In animal dormancy, diapause is the delay in development in response to regular and recurring periods of adverse environmental conditions.Tauber, M.J., Tauber, C.A., Masaki, S. (1986) ''Seasonal Adaptations of Insects''. Oxford University Press It ...
d adults. Most second generation adults diapause in early July. Butterflies of the first generation, reared in captivity in the Emei mountain study, completed their life cycle in 45 to 54 days, with eggs taking 4 to 6 days, caterpillars 21 to 36 days and pupation 10 to 15 days. The breeding took place in temperatures between and humidity of 63.2% to 84.7% on average. Investigations in an artificial climate chamber reveal that
photoperiod Photoperiodism is the physiological reaction of organisms to the length of night or a dark period. It occurs in plants and animals. Plant photoperiodism can also be defined as the developmental responses of plants to the relative lengths of light a ...
and temperature play a role in the larval development and survival rate of the larvae of ''K. inachus''. Photoperiods affect the development period of larvae at but not at and . As temperature increased from 20 °C to 25 °C and 30 °C, the developmental periods of larvae reduced under the same photoperiod to 31.7 to 36.0 days, 26.37 to 27.4 days and 21.0 to 21.5 days, respectively. Increasing temperature also made an increase in the survival rate under different photoperiods. The survival rate of larvae at 20 °C, 25 °C, and 30 °C was 80%–92%, 75%–95%, and 55%–85%, respectively. The low survival rate at 30 °C under most photoperiod gradients requires that artificial breeding of ''K. inachus'' be done below this temperature.


Food plants

The larvae are
polyphagous Feeding is the process by which organisms, typically animals, obtain food. Terminology often uses either the suffixes -vore, -vory, or -vorous from Latin ''vorare'', meaning "to devour", or -phage, -phagy, or -phagous from Greek φαγε ...
, feeding on plants from many families. The list of food plants include: *
Urticaceae The Urticaceae are a family, the nettle family, of flowering plants. The family name comes from the genus ''Urtica''. The Urticaceae include a number of well-known and useful plants, including nettles in the genus ''Urtica'', ramie (''Boehmeri ...
– ''
Girardinia diversifolia ''Girardinia diversifolia'', commonly known as the Himalayan nettle or Nilghiri nettle, is a plant species native to Nepal and in the Himalayan parts of India such as Uttarakhand, Himachal Pradesh and Jammu and Kashmir and in vast parts of China ...
'' in India *
Polygonaceae The Polygonaceae are a family of flowering plants known informally as the knotweed family or smartweed—buckwheat family in the United States. The name is based on the genus ''Polygonum'', and was first used by Antoine Laurent de Jussieu in 1789 ...
– '' Polygonum orientale'' in India *
Rosaceae Rosaceae (), the rose family, is a medium-sized family of flowering plants that includes 4,828 known species in 91 genera. The name is derived from the type genus ''Rosa''. Among the most species-rich genera are ''Alchemilla'' (270), ''Sorbus ...
– ''
Prunus persica ''Prunus'' is a genus of trees and shrubs, which includes (among many others) the fruits plums, cherries, peaches, nectarines, apricots, and almonds. Native to the North American temperate regions, the neotropics of South America, and the paleo ...
'' in India *
Acanthaceae Acanthaceae is a family (the acanthus family) of dicotyledonous flowering plants containing almost 250 genera and about 2500 species. Most are tropical herbs, shrubs, or twining vines; some are epiphytes. Only a few species are distributed in te ...
: ** '' Dicliptera chinensis'' in China ** ''
Hygrophila salicifolia Hygrophila may refer to: * Hygrophila (gastropod), a clade situated under the class and above the superfamily in the gastropods taxonomy * ''Hygrophila'' (plant), a plant genus * '' Astercantha longifolia'', commonly known as hygrophila in En ...
'' in Japan ** ''
Lepidagathis formosensis ''Lepidagathis'' is a genus of plants in the family Acanthaceae. Selected species * ''Lepidagathis ananthapuramensis'' * ''Lepidagathis backeri'' * ''Lepidagathis calycina'' * ''Lepidagathis cristata'' * ''Lepidagathis cuspidata'' * ''Lepidaga ...
'' in Taiwan ** '' Ruellia capitataus'' (syn. ''Strobilanthes capitatus'' & ''S. pentastemonoides'') in India and Taiwan ** '' Rostellularia pracumbens'' in China ** ''
Strobilanthes ''Strobilanthes'' is a genus of about 350 species of flowering plants in the family Acanthaceae, mostly native to tropical Asia and Madagascar, but with a few species extending north into temperate regions of Asia. Many species are cultivated for ...
'' species, in China and Japan, including: ***'' S. flaccidifolius'' in Japan ***'' S. glandulifera'' in Japan ***'' S. tashiroi'' in Japan


Egg

In captive breeding in a net garden, females have been recorded to lay 245.7 eggs on average. This has been increased to 279.8 eggs per female by supplementing nutrition.


Larva

The late stage caterpillar is velvety black, covered with rather long yellowish hair. It has a large number of reddish spines; eleven on each segment, with one dorsal, two subdorsal and three lateral on each side.Based on G. C. Dudgeon as quoted by
Frederic Moore Frederic Moore FZS (13 May 1830 – 10 May 1907) was a British entomologist and illustrator. He produced six volumes of ''Lepidoptera Indica'' and a catalogue of the birds in the collection of the East India Company. It has been said that Mo ...
in Bingham (1905).


Pupa

The pupa is simple. It has a gently keeled thorax. The abdomen displays a series of small conical points dorsally. The colour is light brownish and the pupa is embellished with slaty
irroration This glossary of entomology describes terms used in the formal study of insect species by entomologists. A–C A synthetic chlorinated hydrocarbon insecticide, toxic to vertebrates. Though i ...
s.


Research

The butterfly is considered to be rare in China and consequently much research in its captive breeding has been done. The mitochondrial DNA has been sequenced and found to be 15,183 base pair in size. In addition, the butterfly has been a subject in research on diapause.


See also

*''
Kallima paralekta ''Kallima paralekta'', the Indian leafwing or Malayan leafwing, is a species of Nymphalidae, brush-footed butterfly of the genus ''Kallima''. Despite its common names, it is not found in India or Malaysia, but is endemic to Java and Sumatra of In ...
''


References

{{Taxonbar, from=Q708571 Kallimini Fauna of Pakistan Butterflies of Asia Butterflies described in 1846 Taxa named by Louis Michel François Doyère