Mahasena Corbetti
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''Mahasena corbetti'', the coconut case caterpillar, is a
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 ...
species of
bagworm The Psychidae (bagworm moths, also simply bagworms or bagmoths) are a family of the Lepidoptera (butterflies and moths). The bagworm family is fairly small, with about 1,350 species described. Bagworm species are found globally, with some, su ...
. This species is classified as leaf-eating pest
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 Sym ...
s that produce tough
silk Silk is a natural protein fiber, some forms of which can be woven into textiles. The protein fiber of silk is composed mainly of fibroin and is produced by certain insect larvae to form cocoons. The best-known silk is obtained from the coc ...
out of their bags from dried foliage. Currently, ''M. corbetti'' is considered a 'pest of quarantine importance' in multiple countries and is included on the list of dangerous pests in the
Malaysia Malaysia ( ; ) is a country in Southeast Asia. The federation, federal constitutional monarchy consists of States and federal territories of Malaysia, thirteen states and three federal territories, separated by the South China Sea into two r ...
n plant quarantine act.


Distribution

Although initially native to Malaysia, this destructive leaf defoliating pest has now been recorded in many
Southeast 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 United Nations geoscheme for Asia#South-eastern Asia, south-eastern region of Asia, consistin ...
n regions and
Pacific islands Collectively called the Pacific Islands, the islands in the Pacific Ocean are further categorized into three major island groups: Melanesia, Micronesia, and Polynesia. Depending on the context, the term ''Pacific Islands'' may refer to one of se ...
, including
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 ...
,
Peninsular Malaysia Peninsular Malaysia ( ms, Semenanjung Malaysia; Jawi: سمننجڠ مليسيا), or the States of Malaya ( ms, Negeri-negeri Tanah Melayu; Jawi: نڬري-نڬري تانه ملايو), also known as West Malaysia or the Malaysian Peninsula, ...
,
Sumatra Sumatra is one of the Sunda Islands of western Indonesia. It is the largest island that is fully within Indonesian territory, as well as the sixth-largest island in the world at 473,481 km2 (182,812 mi.2), not including adjacent i ...
,
Java Java (; id, Jawa, ; jv, ꦗꦮ; su, ) is one of the Greater Sunda Islands in Indonesia. It is bordered by the Indian Ocean to the south and the Java Sea to the north. With a population of 151.6 million people, Java is the world's List ...
,
Borneo Borneo (; id, Kalimantan) is the third-largest island in the world and the largest in Asia. At the geographic centre of Maritime Southeast Asia, in relation to major Indonesian islands, it is located north of Java, west of Sulawesi, and eas ...
(Brunei, Sabah),
Solomon Islands Solomon Islands is an island country consisting of six major islands and over 900 smaller islands in Oceania, to the east of Papua New Guinea and north-west of Vanuatu. It has a land area of , and a population of approx. 700,000. Its capita ...
,
Samoa Samoa, officially the Independent State of Samoa; sm, Sāmoa, and until 1997 known as Western Samoa, is a Polynesian island country consisting of two main islands (Savai'i and Upolu); two smaller, inhabited islands (Manono Island, Manono an ...
,
Papua New Guinea Papua New Guinea (abbreviated PNG; , ; tpi, Papua Niugini; ho, Papua Niu Gini), officially the Independent State of Papua New Guinea ( tpi, Independen Stet bilong Papua Niugini; ho, Independen Stet bilong Papua Niu Gini), is a country i ...
and the
Philippines The Philippines (; fil, Pilipinas, links=no), officially the Republic of the Philippines ( fil, Republika ng Pilipinas, links=no), * bik, Republika kan Filipinas * ceb, Republika sa Pilipinas * cbk, República de Filipinas * hil, Republ ...
. As of 2015-2016, it was identified for the first time in a region bordering
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 ...
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 ...
as a pest of areca nut (''Areca catechu'' L.) plants. Because 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. The ...
e of ''M. corbetti'' are most active as pests during bright sunlight, this species thrives in areas that get a lot of natural sunlight and heat.


Morphology

Male ''M. corbetti'' larvae (without the case) are approximately 15-20 mm long. They have a brown thorax and head with a light yellowish brown abdomen. Their cases (or bags) are considerably smaller than that of their female counterparts. Gender determination of the bagworm is difficult in the larval stage, but size distinction in the pupal case lets us approximately determine the sex of the pest before it reaches the adult stage, since females generally have larger cases than males. In the pupal stage, the male pupa are approximately 12-15 mm in length and 3-4 mm in diameter while the female pupa are approximately 20-25 mm in length and 10-12 mm in diameter. This pupa is messy looking in appearance and is made from randomly shaped leaf pieces, stalks and other natural materials. During the pupae stage, male ''M. corbetti'' undergo metamorphosis and turn into moths. These male moths are generally varying degrees of black to brown with white scales on some. They have thinly scaled and often transparent wings and prominently pectinate antennae. Adult females - who spend the entirety of their lives in the pupa - are apterous, cylindrical, and creamy-white with either no or greatly minimized appendages. Both males and females lack
mouthparts Mouthparts may refer to: * The parts of a mouth ** Arthropod mouthparts *** Insect mouthparts {{disambig ...
. The male
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 their ...
being a moth with eyes, antenna, and wings (wingspan of about 22-26 mm) marks
sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most ani ...
in ''M. corbetti''. Anatomically, only adult male individuals of the ''M. corbetti'' species can be
taxon 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 ...
omically identified. However, in this species, there are much fewer males than females and a male takes about 4-5 months to emerge after fertilization, which is why the DNA barcode is more commonly used for taxonomic identification. This DNA barcode was determined by sequencing a standard barcoding region of the mitochondrial oxidase I (COI) gene.


Life cycle

The entire life cycle of ''M. corbetti'' is 110-140 days with a stage by stage breakup as follows: *Egg:10 to 16 days *Larva: 60 to 95 days *Pupa: 14 to 26 days *Adult: 7 to 10 days (male) Neonatal larvae have well developed sclerotized head and thoracic legs as well as weak abdominal forelegs. After emergence from the egg, they partake in an initial dispersal event. This dispersal event has been recorded but not well understood for this species. All that is known is that ''M. corbetti'' larvae are dispersed by wind, vehicles, animals, or humans. The first thing the larvae do once they land on their host is construct their case from nearby foliage. This case is integral to their survival, as it is seen that if larvae are removed from the case, they usually do not construct a new case and will end up dying. This case provides protection, anchorage, and a
microclimate A microclimate (or micro-climate) is a local set of atmospheric conditions that differ from those in the surrounding areas, often with a slight difference but sometimes with a substantial one. The term may refer to areas as small as a few squ ...
environment integral to their survival. If attacked by predators, this case can be lengthened into a sleeve that provides absolute protection for the larvae. The larvae then attach to their host via a strong silk thread, walk around their host, and begin to scrape the surface of the leaves of their host. Whenever the larvae of ''M. corbetti'' are resting, feeding, or mounting, they will ensure that the case is anchored to the leaf via a tough silk thread to prevent falls/contact with the ground. When ready to undergo pupation, the larva attach their case using strong silk thread to the vegetation they were feeding on and enclose the case with more silk thread before positioning themselves with their head towards the posterior end of the case. It is during pupation when dimorphism occurs, with the females barely undergoing
metamorphosis Metamorphosis is a biological process by which an animal physically develops including birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation. Some inse ...
and coming out with a roughly worm-like appearance and the males undergoing significant metamorphosis into winged moths. Once the males emerge, they immediately leave their pupal case. However, females are wingless and stay in their case, adopting a relatively sedentary lifestyle. This has resulted in them evolving degenerate eyes, antennae, and legs in favor of an enlarged egg-sac. It has also resulted in the gender identification of ''M. corbetti'' to be limited to male characteristics.


Mating and reproduction

Since the female resides inside her case,
oviposition The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typical ...
occurs in the interior of the case. Females release pheromones to attract males. Males who want to mate with the female will assemble outside her case and extend his extensive abdomen (which can expand to twice its size during mating) into the female's pupal exuviae. The completion of mating is marked by the female's subsequent oviposition in the pupal skin, resulting in her shrinking to a blob-like mass that falls out of the case. This falling out creates an opening in the posterior of the case through which neonatal larvae crawl out from with the anchorage of a silken thread in some cases. Females have high reproductive potential (with a fecundity measure of over 3000 eggs/female) and in combination with their relatively long time to complete the larval period (about 12-17 weeks in a 5 month life cycle), ''M. corbetti'' are efficient at creating outbreak situations.


Hosts

As a polyphagous pest species, the larvae of ''M. corbetti'' attack the leaves of plants of about 37 genera in 20 families, with their preferred host families being
Fabaceae The Fabaceae or Leguminosae,International Code of Nomenc ...
(to which legume species belong) and
Arecaceae The Arecaceae is a family of perennial flowering plants in the monocot order Arecales. Their growth form can be climbers, shrubs, tree-like and stemless plants, all commonly known as palms. Those having a tree-like form are called palm trees ...
(to which the oil palm species belong). The larva of ''M. corbetti'' attack the leaves in an aggressively destructive manner and the resulting defoliation causes major crop losses, often on economically important plants. Once the larvae start eating the leaves, the leaves turn yellowish in color and fall down upon their death. During the larva stage, the early
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 of ''M. corbetti''  use their
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
s to scrape on the surface of the leaf. This produces holes which cause the leaf to change color and fall prematurely. The leaf tissue also undergoes
necrosis Necrosis () is a form of cell injury which results in the premature death of cells in living tissue by autolysis. Necrosis is caused by factors external to the cell or tissue, such as infection, or trauma which result in the unregulated dige ...
and eventual skeletonisation as a result of this scraping. As the instars mature and become larger, they start cutting the leaf and eventually start chewing the leaf from the margins, causing intense scarring and damage to the leaves. These larvae feed the most in bright sunlight, so they eat most actively in the day.


Parasitoids, predators, and local enemies

The local population size of ''M. corbetti'' is controlled by natural enemies, which vary from palm to palm. In East Malaysia, six species of
hymenopterous Hymenoptera is a large order of insects, comprising the sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are parasitic. Females typic ...
parasitoids (including
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 ...
, Ichneumonidae, and
Eulophidae The Eulophidae are a large family of hymenopteran insects, with over 4,300 described species in some 300 genera. The family includes the genus ''Elasmus'', which used to be treated as a separate family, "Elasmidae", and is now treated as a subf ...
) and five species of
dipterous Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced ...
parasitoids (all Tachinidae) are natural predators of this species. In Singapore, the cleric beetle (''Callimeris bellus'') is a larval predator of ''M. corbetti''. In a study done on the ''M. corbetti'' population in the region bordering India and Bangladesh, two parasitoid species were identified to be naturally parasitizing the ''M. corbetti'' larvae. These species were identified as an unidentified tachinid fly from the order Diptera and an Ichneumon wasp (''
Enteromorpha The sea lettuces comprise the genus ''Ulva'', a group of edible green algae that is widely distributed along the coasts of the world's oceans. The type species within the genus ''Ulva'' is ''Ulva lactuca'', ''lactuca'' being Latin for "lettuce ...
'' spp.). Although the first reports of ''M. corbetti'' in this area were in 2015-2016, this may be inaccurate. It is hypothesized that this species may have entered this area earlier but was unable to attain pest status due to the high parasitoid levels here.


Cultural/economic impact

''Mahasena corbetti'' infestations in Malaysia have accrued a crop loss in excess of 40-50% in the span of two years, resulting in severe economic losses. This is because a moderate defoliation of 10-30% can result in 33-40% crop loss. Although Bt sprays are the current mode of treatment for ''M. corbetti'' control, chemical treatments ineffective after bagworm larvae have metamorphosed to the pupal stage.


References


Further reading

* {{Taxonbar, from=Q13482332 Psychidae Invasive insect species Moths described in 1928