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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 ...
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 16 ...
from
India India, officially the Republic of India, is a country in South Asia. It is the List of countries and dependencies by area, seventh-largest country by area; the List of countries by population (United Nations), most populous country since ...
to
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 ...
. 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 b ...
.


Description

The butterfly wings are shaped like a leaf when in the closed position. When the wings are closed, only the cryptic 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 Masquerades may refer to: Books * ''Masquerade'' (book), a 1979 children's book by Kit Williams that sparked a worldwide treasure hunt * ''Masquerades'' (novel), a 1995 Forgotten Realms novel by Kate Novak and Jeff Grubb * ''Masq ...
, 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 classical architecture, it is a feature of Byzantine and Neoclassical architecture. A horizontal oculus in the center of a dome is also called opaion (; ...
, 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 Municipalities of Switzerland, municipality in the district of Brig (district), Brig in the Cantons of Switzerland, canton of Valais in Switzerland. History Termen is first mentioned in 1201 as ''Terman''. Geography Termen has an ar ...
. 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-Bomb Abomination Absorbing Man Abraxas Abyss Abyss is the name of two characters appearing in Ameri ...
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 hindwi ...
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 or ocellus (sometimes called a pigment pit) is a form of eye or an optical arrangement which has a single lens without the sort of elaborate retina that occurs in most vertebrates. These eyes are called "simple" to distinguish the ...
, 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:Open wing basking position of Kallima inachus (Doyère, (1840)) - Orange Oakleaf WLB.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 () 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 tea plantation, one of ...


Distribution

The orange oakleaf is found in India, Nepal, Bhutan, Bangladesh, and Myanmar, down to
Tenasserim Hills The Tenasserim Hills or Tenasserim Range (, ; , , ; ) is the geographical name of a roughly 1,700 km long mountain chain, part of the Indo-Malayan mountain system in Southeast Asia. Despite their relatively scant altitude these mo ...
. In Southeast Asia it occurs in southern China, Thailand, Laos, Taiwan, and Vietnam. 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 Earth's highest peaks, including the highest, Mount Everest. More than list of h ...
at low elevations, from
Jammu and Kashmir Jammu and Kashmir may refer to: * Jammu and Kashmir (union territory), a region administered by India as a union territory since 2019 * Jammu and Kashmir (state), a region administered by India as a state from 1952 to 2019 * Jammu and Kashmir (prin ...
, 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 * Ga ...
and Kumaon to
West Bengal West Bengal (; Bengali language, Bengali: , , abbr. WB) is a States and union territories of India, state in the East India, eastern portion of India. It is situated along the Bay of Bengal, along with a population of over 91 million inhabi ...
,
Sikkim Sikkim ( ; ) is a States and union territories of India, state in northeastern India. It borders the Tibet Autonomous Region of China in the north and northeast, Bhutan in the east, Koshi Province of Nepal in the west, and West Bengal in the ...
,
Arunachal Pradesh Arunachal Pradesh (; ) is a States and union territories of India, state in northeast India. It was formed from the North-East Frontier Agency (NEFA) region, and India declared it as a state on 20 February 1987. Itanagar is its capital and la ...
, and other states of the northeast. It is also found in central and peninsular India; it flies in
Madhya Pradesh Madhya Pradesh (; ; ) is a state in central India. Its capital is Bhopal and the largest city is Indore, Indore. Other major cities includes Gwalior, Jabalpur, and Sagar, Madhya Pradesh, Sagar. Madhya Pradesh is the List of states and union te ...
,
Jharkhand Jharkhand (; ) is a States and union territories of India, state in East India, 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 ...
,
Chhattisgarh Chhattisgarh (; ) is a landlocked States and union territories of India, state in Central India. It is the List of states and union territories of India by area, ninth largest state by area, and with a population of roughly 30 million, the List ...
,
Orissa Odisha (), formerly Orissa ( the official name until 2011), is a state located in Eastern India. It is the eighth-largest state by area, and the eleventh-largest by population, with over 41 million inhabitants. The state also has the thir ...
and
Andhra Pradesh Andhra Pradesh (ISO 15919, ISO: , , AP) is a States and union territories of India, state on the East Coast of India, east coast of southern India. It is the List of states and union territories of India by area, seventh-largest state and th ...
; i.e. along the central Indian highlands to
Pachmarhi Pachmarhi is a hill station in the Hoshangabad district of Madhya Pradesh state of central India. It has been the location of a cantonment (Pachmarhi Cantonment) since the British Raj. The municipality is located in a valley of the Satpura Rang ...
and
Amarkantak Amarkantak ( NLK ''Amarakaṇṭaka'') is a pilgrim town and a Nagar Panchayat in Anuppur, Madhya Pradesh, India. The Amarkantak region is a unique natural heritage area. It is the meeting point of the Vindhya and the Satpura Ranges, with the ...
, the Western Ghats south to
Bhimashankar Bhimashankar Temple is a Hindu temple dedicated to ''Shiva'' situated in its eponymous village, Bhimashankar, in Pune district of Maharashtra. It is a key pilgrimage centre and contains one of the 12 ''Jyotirlingas''. The temple's Shiva lingam ...
, and in the
Eastern Ghats The Eastern Ghats is a mountain range that stretches along the East Coast of India, eastern coast of the Indian peninsula. Covering an area of , it traverses the states and union territories of India, states of Odisha, Telangana, Andhra Prade ...
north of the river
Godavari The Godavari (, od̪aːʋəɾiː is India's second longest river after the Ganga River and drains the third largest basin in India, covering about 10% of India's total geographical area. Its source is in Trimbakeshwar, Nashik, Maharash ...
. 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 (occasionally unhyphenated; 15 August 1906 – 16 April 1963) 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 ...
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 ChongqingPostal Romanization, Previously romanized as Chungking ();. is a direct-administered municipality in Southwestern China. Chongqing is one of the four direct-administered municipalities under the State Council of the People's Republi ...
municipality,
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
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 Puddling is a behaviour in which an organism seeks out nutrients in certain moist substances such as rotting plant matter, mud, and carrion, and sucks up the fluid. Where the conditions are suitable, conspicuous insects such as butterflies common ...
. 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 (20 ...
'' species), and some bacteria.


Life cycle

In the Himalayas, the butterfly is multivoltine 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, which occurs between each moult (''ecdysis'') until sexual maturity is reached. Arthropods must shed the exoskeleton in order to ...
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 Mount Emei (; zh, c=峨眉山, p=Éméi shān, O2-mei2 shan1), alternatively Mount Omei, is a mountain in Sichuan Province, China, and is the highest of the Four Sacred Buddhist Mountains of China. Mount Emei sits at the western rim of th ...
(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 Photoperiod is the change of day length around the seasons. The rotation of the earth around its axis produces 24 hour changes in light (day) and dark (night) cycles on earth. The length of the light and dark in each phase varies across the season ...
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 (''Boehmeria ...
– ''
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 (biology), family of flowering plants known Common name, informally as the knotweed family or smartweed—buckwheat family in the United States. The Botanical name, name is Basionym, based on the genus ''Polygonum'', ...
– '' Polygonum orientale'' in India *
Rosaceae Rosaceae (), the rose family, is a family of flowering plants that includes 4,828 known species in 91 genera. The name is derived from the type genus '' Rosa''. The family includes herbs, shrubs, and trees. Most species are deciduous, but som ...
– ''
Prunus persica The peach (''Prunus persica'') is a deciduous tree first domesticated and cultivated in China. It bears edible juicy fruits with various characteristics, most called peaches and the glossy-skinned, non-fuzzy varieties called nectarines. Peach ...
'' in India *
Acanthaceae Acanthaceae () is a Family (biology), family (the acanthus family) of dicotyledonous flowering plants containing almost 250 genera and about 2500 species. Most are Tropics, tropical Herbaceous plant, herbs, shrubs, or twining vines; some are epip ...
: ** ''
Dicliptera chinensis ''Dicliptera'' is a genus of flowering plants in the Acanthus (plant), bear's breeches family, Acanthaceae. It includes 223 species native to the tropics and subtropics worldwide. Well-known synonyms include ''Peristrophe'' and ''Dactylostegium' ...
'' in China ** ''
Hygrophila salicifolia Hygrophila may refer to: * Hygrophila (gastropod) Hygrophila is a taxonomic superorder of air-breathing freshwater snails, aquatic pulmonate gastropod mollusks within the clade Panpulmonata.MolluscaBase eds. (2020). MolluscaBase. Hygrophila ...
'' in Japan ** '' Lepidagathis formosensis'' 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 (biology), family Acanthaceae, mostly native plant, native to tropical Asia and Madagascar, but with a few species extending north into temperateness, temperate r ...
'' 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 irrorations.


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 in In ...
''


References

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