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''Hyblaea puera'', the teak defoliator, is a
moth Moths are a paraphyletic group of insects that includes all members of the order Lepidoptera that are not butterflies, with moths making up the vast majority of the order. There are thought to be approximately 160,000 species of moth, many of w ...
and
cryptic species complex In biology, a species complex is a group of closely related organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The taxa in the complex may be able to hybridize readily with each oth ...
native to
South Asia South Asia is the southern subregion of Asia, which is defined in both geographical Geography (from Greek: , ''geographia''. Combination of Greek words ‘Geo’ (The Earth) and ‘Graphien’ (to describe), literally "earth descr ...
and
South-east Asia Southeast Asia, also spelled South East Asia and South-East Asia, and also known as Southeastern Asia, South-eastern Asia or SEA, is the geographical south-eastern region of Asia, consisting of the regions that are situated south of mainland ...
. It was first described by
Pieter Cramer Pieter Cramer (21 May 1721 (baptized) – 28 September 1776), was a wealthy Dutch merchant in linen and Spanish wool, remembered as an entomologist. Cramer was the director of the Zealand Society, a scientific society located in Flushing, and a mem ...
in 1777. The species has also been recently reported to be present in
Central America Central America ( es, América Central or ) is a subregion of the Americas. Its boundaries are defined as bordering the United States to the north, Colombia to the south, the Caribbean Sea to the east, and the Pacific Ocean to the west. ...
and Africa. The caterpillar feeds on
teak Teak (''Tectona grandis'') is a tropical hardwood tree species in the family Lamiaceae. It is a large, deciduous tree that occurs in mixed hardwood forests. ''Tectona grandis'' has small, fragrant white flowers arranged in dense clusters ( pan ...
and other trees. It is considered to be one of the major teak pests around the world.


Distribution and habitat

''Hyblaea puera'' occurs in forests across southern Asia 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 ...
, Sri Lanka and
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 ...
, through
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 b ...
and the rest of south-east Asia to
New Guinea New Guinea (; Hiri Motu: ''Niu Gini''; id, Papua, or , historically ) is the world's second-largest island with an area of . Located in Oceania in the southwestern Pacific Ocean, the island is separated from Australia by the wide Torr ...
, and north
Queensland ) , nickname = Sunshine State , image_map = Queensland in Australia.svg , map_caption = Location of Queensland in Australia , subdivision_type = Country , subdivision_name = Australia , established_title = Before federation , establishe ...
in Australia. Recent reports place it in Guadeloupe, and Suriname.


Life cycle

The adult moths are comparatively small, with a wingspan of 3–4 cm, and have a characteristic resting posture that conceals the black and orange-yellow hindwings under the greyish-brown forewings. Males and females emerge more or less simultaneously and mating takes place within a couple of days. Eggs are laid on tender new leaves, placed singly near the veins, and usually on the undersurface. They are oval, flat, and white and measure about 1 mm in length. About 500 eggs are laid per female with a recorded maximum of 1000. Larvae hatch in about two days. There are five larval
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 ...
s. The first and second instars mainly feed on the leaf surface. Starting with the third instar, the larva cuts out a leaf flap, usually at the edge of the leaf, folds it over, fastens it with silk, and feeds from within. The entire leaf, excluding the major veins of tender leaves, is eaten, but more veins are left in older leaves. Under the optimal conditions, the larval period lasts 10 to 12 days. The full-grown larva measures about 3.5–4.5 cm, and there is considerable colour variation in the fourth and fifth instars; the body may be either wholly black or dark greyish to black, with longitudinal coloured bands which may include a dorsal orange or ocherous band and lateral white lines. A recent study revealed the existence of density-dependent colour polyphenism and resistance build-up against invading baculovirus by ''H. puera'' larvae. The mature larvae descend to the ground on silken threads and pupate under a thin layer of leaf litter or soil, within loosely built cocoons made of dry or decayed leaves, or soil particles held together with silk. Pupation may some times occur within green leaves of other plants in the under growth, folded or juxtaposed with silk. On ''
Avicennia ''Avicennia'' is a genus of flowering plants currently placed in the bear's breeches family, Acanthaceae. It contains mangrove trees, which occur in the intertidal zones of estuarine areas and are characterized by its "pencil roots", which are ...
'' plants the caterpillars pupate inside the cases made out of the host plant leaves themselves. It is probably an adaptive trait acquired by the species for surviving in the hostile mangrove environment, since pupation is not possible in the muddy and inundated soils of mangroves. The average pupal period lasts six to eight days under optimal conditions. There is no evidence of hibernation or aestivation of the pupa. File:Hyblaea puera larva (29797907483).jpg, Larva File:Hyblaea puera (The Teak Defoliator) (2) (29798819784).jpg, Pupa


Temporal and spatial dynamics

The teak defoliator is present year round in teak plantations, but in varying population densities. During the period of natural defoliation of teak (November, December, and January), the pest density is very low (endemic). Every year high-intensity outbreaks of teak defoliator occur immediately after the premonsoon showers in late February or early march in
Kerala Kerala ( ; ) is a state on the Malabar Coast of India. It was formed on 1 November 1956, following the passage of the States Reorganisation Act, by combining Malayalam-speaking regions of the erstwhile regions of Cochin, Malabar, South ...
. These centres are highly localized outbreaks which represent the transitional stage between very sparse endemic population and high density outbreak population. These centres will be 5,000 to 15,000 square metres in area and are characterized by heavy tree top infestation. The months of April, May, June, and July witness a series of large outbreaks. During late July or September, the population declines to the endemic level. In some years, there will be fresh outbreaks during October. From then on until the next year, the population remains at the endemic level.


Natural control


Parasitoids

The main parasitoids attacking teak defoliator include the
tachinid The Tachinidae are a large and variable family of true flies within the insect order 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 this family ...
'' Palexorisa solennis'', a eulophid '' Sympiesis hyblaeae'', the chalicid '' Brachymeria lasus'' and the three ichneumonids, ''Eriborus gardneri'', ''Stictopisthus'' sp. and ''Echthromorpha agrestoria notulatoria''. ''B. lasus'' is a pupal parasitoid and all others are larval parasitoids. The overall parasitism by all species is about 9%.


Predators

Wasps, spiders, birds, and
bonnet macaque The bonnet macaque (''Macaca radiata''), also known as zati,Chambers English Dictionary is a species of macaque endemic to southern India. Its distribution is limited by the Indian Ocean on three sides and the Godavari and Tapti Rivers, along wit ...
s eat ''H. puera''. Forty-eight species of birds have been recorded as feeding on teak defoliator larvae during large-scale outbreaks.


Pathogens

The bacteria ''
Enterobacter aerogenes ''Klebsiella aerogenes'', previously known as ''Enterobacter aerogenes,'' is a Gram-negative, oxidase negative, catalase positive, citrate positive, indole negative, rod-shaped bacterium. The bacterium is approximately 1-3 microns in length, an ...
'', '' Bacillus thuringiensis'', ''
Pseudomonas aeruginosa ''Pseudomonas aeruginosa'' is a common encapsulated, gram-negative, aerobic–facultatively anaerobic, rod-shaped bacterium that can cause disease in plants and animals, including humans. A species of considerable medical importance, ''P. aerug ...
'' and ''
Serratia marcescens ''Serratia marcescens'' () is a species of rod-shaped, Gram-negative bacteria in the family Yersiniaceae. It is a facultative anaerobe and an opportunistic pathogen in humans. It was discovered in 1819 by Bartolomeo Bizio in Padua, Italy.Serra ...
'' are identified as causing mortality to the teak defoliator. A synnematous fungus of the genus ''
Hirsutella ''Hirsutella'' is a genus of asexually reproducing fungi in the Ophiocordycipitaceae family. Originally described by French mycologist Narcisse Théophile Patouillard in 1892, this genus includes species that are pathogens of insects, mites and ne ...
'' is found to be pathogenic to this pest. An absolutely specific virus with refractile polyhedral inclusion bodies, staining blue in
Giemsa Giemsa stain (), named after German chemist and bacteriologist Gustav Giemsa, is a nucleic acid stain used in cytogenetics and for the histopathological diagnosis of malaria and other parasites. Uses It is specific for the phosphate groups of ...
and thick blue in Buffalo Black, named as ''Hyblaea puera'' nucleopolyhedrovirus (HpNPV) is found to be very effective in the biological control of this pest.


''Hyblaea puera'' nucleopolyhedrovirus (HpNPV)

Large-scale deaths of teak defoliator larvae characterized by cessation of feeding, flaccidity and subsequent liquefaction of body tissues have been reported by Stebbing as early as 1903. During 1985, an investigation of microbial pathogens of ''H. puera'' was undertaken in the plantations of
Nilambur Nilambur is a major town, a municipality and a Taluk in the Malappuram district of the Indian state of Kerala. It is located close to the Nilgiris range of the Western Ghats on the banks of the Chaliyar River. This place is also known as 'Teak ...
Forest Divisions of Kerala, India, by KFRI detected several dead insects with the characteristic symptoms as observed by Stebbing. Microscopic observation of tissues revealed the presence of refractile polyhedral inclusions bodies, which stained blue in Giemsa, measured 0.9–2.4 micrometers in diameter in the scanning electron micrograph taken by Jean Adams at USDA, confirmed its identity as NPV. The NPVs come under the family of baculoviridae and its virions are enveloped rod shaped nucleocapsids containing circular, supercoiled, double stranded DNA.


Mode of action

The NPV which enters the insect gut, lyses in the alkaline environment of the midgut, releasing virions. Virions invade the columnar cells of midgut epithelium and integrate DNA into the nucleus of midgut cells. Later the viral DNA takes over control of the cellular machinery to reproduce itself. The ECVs which are the progenies released into the haemocoel from the midgut cells are more infectious than the PIBs and mediate disease spread within the insect body. A rapid spread of infection in the insect body leads immediately to cessation of feeding and later on, to death. Even in the fully mature larvae, HpNPV can kill in 60–72 hours, making it one of the fastest acting insect viruses. The discovery of virus disease on ''H. puera'' was a major breakthrough in the teak defoliator management research.


As food

The teak moth, in Indonesia known as entung jati, is commonly eaten in the regions where they thrive. They are usually caught for consumption at the cocoon stage, at which point they are easily collected from the ground. This serves as an essential, readily accessible source of protein for the villagers, commonly farmers, around these areas.


References

* Senthil-Nathan, S., Saehoon, K., (2006). Behavioral and physiological effects of ''Melia azedarach'' L. extract on the teak defoliator ''Hyblaea puera'' Cramer (Lepidoptera: Hyblaeidae). ''Crop Protection'' 25 (3), 287-291 (Published by Elsevier Inc. UK). * Nair, K. S. S., Sudheendrakumar, V. V., Varma, R. V., and Chacko, K. C. (1985). Studies on the seasonal incidence of defoliators and the effect of defoliation on volume increment of teak. Research Report, Kerala Forest Research Institute. 30, 78pp. * Nair, K. S. S. and Sudheendrakumar, V. V. (1986). The teak defoliator, ''Hyblaea puera'': Defoliation dynamics and evidence for short – range migration of moths. ''Proceedings of the Indian Academy of Sciences'' (Anima. Sci.) 95(1):7-21. * Sudheendrakumar, V.V. (1986). Studies on natural enemies of the teak pests, ''Hyblaea puera'' and ''Eutectona machaeralis''. Research Report No.38, Kerala Forest Research Institute, Peechi, 23pp. *Sudheendrakumar, V.V. (1990). A report on the Ichneumonid parasitoids of ''Hyblaea puerarecorded'' from Nilambur, Kerala. ''Journal of Tropical Forestry'' 6(1):102-103. * Sajeev TV, 2000. Spatial dynamics of teak defoliator (''Hyblaea puera'' Cramer) outbreaks: Patterns and Causes. PhD Thesis. Cochin University of Science and Technology.101 * Sudheendrakumar, V. V. (1997). Evaluation of parasitoids for the biological control of the teak defoliator. Research report, Kerala Forest Research Institute. 129, 32p. * Zacharias, V. J. and Mohanadas, K. (1990). Bird predators of the teak defoliator, ''Hyblaea puera''. ''Indian Journal of Forestry''. 13, 122-127pp. * Mohammed Ali, M. I. and Sudheendrakumar, V. V. (1991). Possible use of microbial pathogens against teak pests. Proceedings of the International Teak Symposium, Trivandrum. * Sudheendrakumar, V. V., Mohammed Ali, M. I. and Varma, R. V. (1988). Nuclear polyhedrosis virus of the teak defoliator, ''Hyblaea puera''. ''Journal of Invertebrate Pathology''. 51, 307–308. {{Taxonbar , from=Q5952973 Hyblaeidae Moths of Guadeloupe Moths of Japan Moths of Réunion Moths of Africa Moths of the Middle East