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 ...
s that produce tough
silk Silk is a natural fiber, natural protein fiber, some forms of which can be weaving, woven into textiles. The protein fiber of silk is composed mainly of fibroin and is most commonly produced by certain insect larvae to form cocoon (silk), c ...
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. Featuring the Tanjung Piai, southernmost point of continental Eurasia, it is a federation, federal constitutional monarchy consisting of States and federal territories of Malaysia, 13 states and thre ...
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 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 ...
n regions and
Pacific islands The Pacific islands are a group of islands in the Pacific Ocean. They 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 several ...
, including
Thailand Thailand, officially the Kingdom of Thailand and historically known as Siam (the official name until 1939), is a country in Southeast Asia on the Mainland Southeast Asia, Indochinese Peninsula. With a population of almost 66 million, it spa ...
,
Peninsular Malaysia Peninsular Malaysia, historically known as Malaya and also known as West Malaysia or the Malaysian Peninsula, is the western part of Malaysia that comprises the southern part of the Malay Peninsula on Mainland Southeast Asia and the list of isla ...
,
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 list of islands by area, sixth-largest island in the world at 482,286.55 km2 (182,812 mi. ...
,
Java Java is one of the Greater Sunda Islands in Indonesia. It is bordered by the Indian Ocean to the south and the Java Sea (a part of Pacific Ocean) to the north. With a population of 156.9 million people (including Madura) in mid 2024, proje ...
,
Borneo Borneo () is the List of islands by area, third-largest island in the world, with an area of , and population of 23,053,723 (2020 national censuses). Situated at the geographic centre of Maritime Southeast Asia, it is one of the Greater Sunda ...
(Brunei, Sabah),
Solomon Islands Solomon Islands, also known simply as the Solomons,John Prados, ''Islands of Destiny'', Dutton Caliber, 2012, p,20 and passim is an island country consisting of six major islands and over 1000 smaller islands in Melanesia, part of Oceania, t ...
,
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 ...
,
Papua New Guinea Papua New Guinea, officially the Independent State of Papua New Guinea, is an island country in Oceania that comprises the eastern half of the island of New Guinea and offshore islands in Melanesia, a region of the southwestern Pacific Ocean n ...
and the
Philippines The Philippines, officially the Republic of the Philippines, is an Archipelagic state, archipelagic country in Southeast Asia. Located in the western Pacific Ocean, it consists of List of islands of the Philippines, 7,641 islands, with a tot ...
. As of 2015-2016, it was identified for the first time in a region bordering
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 ...
and
Bangladesh Bangladesh, officially the People's Republic of Bangladesh, is a country in South Asia. It is the List of countries and dependencies by population, eighth-most populous country in the world and among the List of countries and dependencies by ...
as a pest of
areca nut The areca nut ( or ) or betel nut () is the fruit of the areca palm (''Areca catechu''). The palm is originally native to the Philippines, but was carried widely through the tropics by the Austronesian migrations and trade since at least 15 ...
(''Areca catechu'' L.) plants. Because the
larva A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
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. The male
pupa A pupa (; : 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 life cycle, the stages th ...
being a moth with eyes, antenna, and wings (wingspan of about 22-26 mm) marks
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 ...
in ''M. corbetti''. Anatomically, only adult male individuals of the ''M. corbetti'' species can be
taxon In biology, a taxon (back-formation from ''taxonomy''; : 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 name and ...
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 DNA barcoding is a method of species identification using a short section of DNA from a specific gene or genes. The premise of DNA barcoding is that by comparison with a reference library of such DNA sections (also called "sequences"), an indivi ...
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 Sclerosis (also sclerosus in the Latin names of a few disorders) is a hardening of tissue and other anatomical features. It may refer to: * Sclerosis (medicine), a hardening of tissue * in zoology, a process which forms sclerites, a hardened exo ...
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 atmosphere of Earth, atmospheric conditions that differ from those in the surrounding areas, often slightly but sometimes substantially. The term may refer to areas as small as a few square m ...
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 transformation or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and different ...
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 typica ...
occurs in the interior of the case. Females release
pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
s 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 In biology, exuviae are the remains of an exoskeleton and related structures that are left after ecdysozoans (including insects, crustaceans and arachnids) have molted. The exuviae of an animal can be important to biologists as they can often be ...
. 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 Fecundity is defined in two ways; in human demography, it is the potential for reproduction of a recorded population as opposed to a sole organism, while in population biology, it is considered similar to fertility, the capability to produc ...
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 Fabaceae () or Leguminosae,International Code of Nomen ...
(to which legume species belong) and
Arecaceae The Arecaceae () is a family (biology), family of perennial plant, perennial, flowering plants in the Monocotyledon, monocot order Arecales. Their growth form can be climbing palm, climbers, shrubs, tree-like and stemless plants, all commonly k ...
(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, which occurs between each moult (''ecdysis'') until sexual maturity is reached. Arthropods must shed the exoskeleton in order to ...
s of ''M. corbetti''  use their
mandible In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...
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. The term "necrosis" came about in the mid-19th century and is commonly attributed to German pathologist Rudolf Virchow, who i ...
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 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 The Ichneumonidae, also known as ichneumon wasps, ichneumonid wasps, ichneumonids, or Darwin wasps, are a family of parasitoid wasps of the insect order Hymenoptera. They are one of the most diverse groups within the Hymenoptera with roughly 25 ...
, and
Eulophidae The Eulophidae is 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 subfa ...
) and five species of dipterous parasitoids (all
Tachinidae The Tachinidae are a large and variable family of true fly, flies within the insect order Fly, 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 t ...
) 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