Tessaratomidae
Encyclopedia
Tessaratomidae is a family
of true bugs
. It contains about 240 species
of large bugs divided into 3 subfamilies and 56 genera
.
Tessaratomids resemble large stink bugs (family Pentatomidae
) and are sometimes quite colorful. Most tessaratomids are Old World
, with only three species known from the Neotropics. Some members of Tessaratomidae exhibit maternal care of eggs and offspring. The defensive chemicals of certain species can cause significant damage if they come into contact with human skin; they may also cause temporary blindness.
All species are exclusively plant-eaters, some of major economic importance as agricultural pests. A few species are also consumed as human food in some countries.
and are referred to mostly as tessaratomids.
Tessaratomids are ovate to elongate-ovate bugs. They range in size from the smallest members of the tribe
Sepinini at 6 to 7 mm (0.236220472440945 to 0.275590551181102 in), to the large Amissus atlas of tribe Tessaratomini at 43 to 45 mm (1.7 to 1.8 in). They are generally quite large and usually exceed 15 mm (0.590551181102362 in) in length.
The head of tessaratomids is generally small and triangular, with the antennae
having 4 to 5 segments (though some of them, for example Siphnus, have relatively large heads). The scutellum (Latin for 'little shield', the hard extension of the thorax
covering the abdomen
in hemiptera
ns) is triangular and does not cover the leathery middle section of the forewing but is often partially covered by the prothorax
. The tarsi (the final segments of the legs) have 2 to 3 segments.
Like all hemiptera
ns, instead of mandibles for chewing, tesseratomids possess a piercing-sucking mouthpart for feeding (known as the rostrum). In tesseratomids, the rostrum has 4 segments.
Tessaratomids are oftentimes vividly colored.
and Sapindales
, and spend most of their lives in tree leaves and stems. They exhibit incomplete metamorphosis and have lifespans that can be several years.
Some tessaratomids guard their eggs and nymphs from predators which may include parasitoid
wasp
s and assassin bugs.
The eggs are laid in compact clusters glued to the leaves of a variety of plants. The laying arrangement can follow a pattern. For example, in Pygoplatys tenangau, the egg clusters are distinctively hexagonal; while in Piezosternum subulatum, they are arranged in two neat rows. The eggs are usually initially white, cream, or yellow in color but can change as the embryo
s inside mature.
Nymphs emerge from the eggs through peristaltic movement
s and with the help of an internal nearly H-shaped structure in the egg known as the 'egg burster'.
As in other hemiptera
ns, tessaratomids are hemimetabolic
, undergoing incomplete metamorphosis
. This means that they do not possess larva
l and pupa
l stages. Instead, juvenile tessaratomids (called nymphs
), hatch directly from the eggs. The nymphs resemble fully grown adults except for size and the absence of wings.
Nymphs usually undergo four to five successive stages of moltings (ecdysis
), increasing in size and becoming more adult-like with each stage until the final molting. The stages are individually known as instar
s, with the earliest stage (just after hatching) being known as the first nymphal instar. Nymphs may also differ significantly from adults in colors and patterns exhibited. In some species, nymphs often exhibit strikingly vibrant colors in contrast to the relative drabness of adults. The colors can also vary between instars.
Mating between adults can last for several hours, with the male and female attached end-to-end.
is a well-documented presocial behavior
among tessaratomids. Egg guarding by adult females were first observed in 1991 by S. Tachikawa among Japanese species of the genera Pygoplatys (subfamily Tessaratominae) and Erga
(subfamily Oncomerinae).
In 1998, Gogala et al. described tessaratomines of the genus Pygoplatys from Thailand
and Malaysia showing egg guarding behavior. In addition, they were also observed to exhibit another remarkable maternal behavior. A dense cluster of small nymphs were photographed being carried around by adult females. The nymphs were firmly clutching unto the bottom side of the abdomens of the adults and to each other, forming a compact mass. The females seem unimpeded by their burden and were able to walk around normally and even fly. The nymphs, however, were not observed feeding. This behavior is known as "nymphal phoresy" (used adjectivally
as "phoretic").
In the Indonesia
n species Pygoplatys tenangau, females will cover the clutch of 70 to 120 eggs with their bodies after laying them, literally "standing guard" over them. When approached, they will spray defensive liquid at perceived attackers and may buzz their wings. They will not willingly abandon the eggs they are guarding, however, and if picked up will try to hold unto the leaf where their eggs are attached. It usually takes slightly more than two weeks for the eggs to hatch. The hatching process will take up 3 to 4 days, during which the newly hatched nymphs will immediately clamber unto their mother's abdomen. They were observed to remain phoretic for at least 17 days (Magnien et al., 2008).
In the subfamily Oncomerinae, a predominantly Australia
n group of large colorful bugs, brooding behavior varies from species which do not practice it at all (exhibited by Musgraveia sulciventris) to adult females carrying first and second instar
nymphs on their abdomens (Monteith, 2011).
Adult female oncomerines of the genus Lyramorpha will guard nymphs at least to the second instar.
Oncomerines of the genera Cumare, Garceus, and Peltocopta exhibit the most advanced form of maternal care. Like the previously described Southeast Asian Pygoplatys individuals, the females actually carry young nymphs around on their abdomens. As the nymphs grow older, they eventually separate from their mothers, lose their bright colors, and become more solitary prior to molting into adults. Species which exhibit this behavior often have significantly flattened and expanded abdomens.
Of the Australian oncomerines, the bronze orange bug (Musgraveia sulciventris) is the only species unequivocally documented to lack maternal brooding behavior. This peculiarity might be connected to the unreliability of the food plant availability for the species (Monteith, 2011). Unlike other oncomerines who can only lay one egg clutch for the certain amount of time it takes to care for them, M. sulciventris can produce multiple egg clutches rapidly because females don't have to care for them. This allows M. sulciventris to rapidly expand their population when conditions are favorable.
s), the source of the common name for pentatomoids - 'stink bugs'. When threatened, tessaratomids may squirt a strong jet of caustic liquid up to a distance of 15 to 27 cm (5.9 to 10.6 in).
The chemicals produced by heteroptera
ns are usually alkane
s and aldehyde
s from gland
s in the thorax
. Compounds that are primarily for protection against fellow arthropod
s (to which they are lethal). However, the defensive chemicals of tessaratomids (particularly that of Tessaratoma papillosa and Musgraveia sulciventris) are notable for being one of the most debilitating to vertebrate
s, probably a defense specifically aimed against birds. They can cause damage to human skin and even cause temporary blindness if sprayed unto the eyes.
In Lyramoprha parens, nymphs are also known to be highly gregarious, forming massed feeding groups and moving to new feeding sites in close-packed groups. This behavior, along with their bright colorations and stink glands is believed to help in discouraging potential predators.
Aggregation behavior is also common in adults in some species. Aside from combined chemical defenses, other possible benefits of aggregation include better mating opportunities and shelter, greater retention of moisture and heat, and a possible sense of security. Lone bugs in some species were known to be more likely to be skittish than bugs in groups. However, aggregation can also increase the threats of disease
s, parasites, and parasitoid
s.
If all these defenses fail, tessaratomids will escape predators either by flying away or dropping to the ground (except in cases of females guarding eggs as discussed above).
of tessaratomids include several tiny parasitoid
wasp
s as well as other hemiptera
ns.
Parasitoid wasps that parasitize tessaratomids usually come from the families Eupelmidae
, Scelionidae
, and Encyrtidae
. Adult female parasitoid wasps will search out eggs laid by tessaratomoids. Upon finding some, they will thrust their ovipositor
s into them and lay eggs inside. The eggs of parasitoid wasps hatch and develop inside the tessaratomid eggs, feeding on the tessaratomid embryo and eventually killing it. Infested eggs characteristically turn darker in color as the wasp larva matures. After about a week, one or more adult wasps will then emerge from the now empty egg.
Musgraveia sulciventris is parasitized by the wasps Eupelmus poggioni and Telenomus spp.; Tessaratoma javanica by the wasps Anastatus colemani and Anastatus kashmirensis(?); and the lychee giant stink bug Tessaratoma papillosa by the wasps Ooencyrtus phongi, Anastatus spp. (particularly Anastatus japonicus), and Trissolcus spp. (particularly Trissolcus latisulcus).
In tessaratomids considered to be agricultural pests (like the Musgraveia sulciventris and Tessaratoma papillosa), the wasps that parasitize them are being studied as potential biological control agents
. In the Fujian
, Guangdong
, and Guangxi
provinces of China, mass-reared Anastatus japonicus are being released to combat Tessaratoma papillosa pests in lychee and longan
crops. The same measures are also reportedly being done in Thailand.
Musgraveia sulciventris is also preyed upon by the predatory pentatomid
Asopus and assassin bugs (family Reduviidae
) of the species Pristhesancus papuensis and Pristhesancus plagipennis.
and among the Venda people
of South Africa
. The insects are light green in color and quite large, averaging at 25 mm (0.984251968503937 in) in length. They are most widely known in South Africa as "thongolifha", though they are also known as "tsonônô". In Zimbabwe, they are known as "harurwa" or "harugwa".
Encosternum delegorguei are collected just before dawn when they are least active and are easier to catch. They are caught carefully, taking care not to kill them. The chemicals released by the bugs can often stain the hands of collectors orange if they collect them barehanded. The bugs which died during collection are carefully separated from live bugs. This is because the chemicals stored in the stink glands are unpalatable - being extremely bitter. As dead bugs can not release the remaining chemicals in their bodies, they are deemed unsuitable for consumption and discarded. The remaining live bugs are placed in a bucket with a small amount of warm water. This is then carefully agitated so as to make them release all their defense chemicals in alarm. This is repeated several more times until their stink glands are drained.
The live bugs with their now empty stink glands are then boiled in water. Further sorting is done afterwards. Dead bugs which died before they could release all their chemicals can be distinguished from the 'clean' bugs by their blackened abdomens after boiling. These are also rejected. The remaining bugs are then dried under the sun.
In cases where the bugs were collected dead, another method is used to remove the chemicals. The bugs are beheaded and carefully squeezed so chemicals in their stink glands flow out the severed neck. The liquids secreted are then wiped off and the bugs boiled and sun-dried like the previous procedure.
After removing the wings, the dried bugs can be eaten as is, fried with a little salt, or cooked with a type of porridge called pap
. They are believed to be a good source of protein
.
Diminishing harvests of E. delegorguei has been a cause for concern in recent years. It may be due to the decline in the number of available food plants which are being harvested locally for firewood. Studies are being done in South Africa for ways to ensure sustainable harvests of E. delegorguei, as well as for the possibilities of rearing them in captivity for human consumption.
In Thailand
(where a total of 81 insect species are reportedly eaten), large tessaratomids of the genera Pygoplatys and Tessaratoma (T. papillosa and T. javanica) are eaten.
In Laos
, Tessaratoma quadrata, locally known as "mien kieng" are also eaten. The same species is also eaten among the Galo people of Northeast India
where they are known as "tari". Only adults are consumed. The wings are removed and the bugs eaten raw or cooked into chutney
.
. They also feed on the closely related Sapindaceae
fruit trees like longan
(Dimocarpus longan) and rambutan
(Nephelium lappaceum). The closely related Tessaratoma quadrata and Tessaratoma javanica are also minor pests of apple
, pear
, and lychee trees.
Bronze orange bugs (Musgraveia sulciventris) are serious pests to citrus
crops in Australia
. They are very large bugs, around 20 mm (0.78740157480315 in) in length, whose native host plants are members of the rue family, Rutaceae
.
In Indonesia
, the tessaratomid Pygoplatys tenangau, locally known as the "tenangau", is considered to be one of the most important pests of Damar gardens. Damar gardens are cultivated forests of trees of the genera Shorea
, Balanocarpus, or Hopea
used as a source of Damar resin
. P. tenangau is the only known tessaratomid which feeds on Dipterocarpaceae
.
In Papua New Guinea
, Agapophyta viridula and Agapophyta similis are regarded as pests of Tephrosia
spp. and pigeon pea
s (Cajanus cajan). Agapophyta bipunctata are known minor pests of coconut
s (Cocos nucifera) and sago
(Metroxylon sagu) as well.
in 1864. In 1900, the Hungarian entomologist Géza Horváth
divided the family into 9 tribes
and established a key to determining genera. The English entomologist George Willis Kirkaldy
increased the number of subfamilies under Tessaratomidae to 11 in 1909. Dennis Leston (1955) followed Kirkaldy's system but reclassified some tribes to subtribes. The current classification is based on the work of Pramod Kumar in 1969 who reduced the number of subfamilies to three - Natalicolinae, Oncomerinae, and Tessaratominae; with Tessaratominae being further divided into three tribes - Prionogastrini, Sepinini and Tessaratomini. Subsequent revisions by Sinclair (1989), Rolston et al. (1993), Schuch & Slater (1995), Sinclair (2000), Cassis & Gross (2002), and Rider (2006), are all based upon Kumar's system.
Tessaratomidae is classified under order
Hemiptera
(true bugs), suborder Heteroptera
, infraorder Pentatomomorpha
, and superfamily
Pentatomoidea (shield bugs and stink bugs). It is currently divided into three subfamilies: Natalicolinae (with 8 genera), Oncomerinae (with 15 genera), and Tessaratominae (with 33 genera and one of uncertain placement).
They are mostly found in tropical Africa, Asia, and Oceania though a few species can be found in the Neotropics and Australia
. There are about 240 species known.
Listed below are the three subfamilies; their authors and type genera; the tribes, subtribes, and genera classified under them; and their distribution ranges
:
of Germany
; but the specimen was published with no formal description and is thus unacceptable as a valid taxon
. Another fossil genus, Latahcoris, from the Miocene
Latah Formation
of Idaho
, was described in 1931 by T.D.A. Cockerell.
Pentatomoidea in 2008 hints that Tessaratomidae and Dinidoridae represented a monophyletic group. However, the difficulty in securing enough materials for examination for both groups leaves this as yet unresolved.
Below is the morphological
unweighted tree
of the superfamily Pentatomoidea after Grazia et al. (2008). Tessaratomidae is in bold. Both Dinidoridae and Tessaratomidae are shown in dotted lines, signifying uncertain status.
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...
of true bugs
Hemiptera
Hemiptera is an order of insects most often known as the true bugs , comprising around 50,000–80,000 species of cicadas, aphids, planthoppers, leafhoppers, shield bugs, and others...
. It contains about 240 species
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...
of large bugs divided into 3 subfamilies and 56 genera
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
.
Tessaratomids resemble large stink bugs (family Pentatomidae
Pentatomidae
Pentatomidae, Greek pente meaning five and tomos meaning section, is a family of insects belonging to order hemiptera that includes some of the stink bugs and shield bugs. Their antennae are 5-segmented, which gives the family its scientific name. Their bodies are usually shield-shaped...
) and are sometimes quite colorful. Most tessaratomids are Old World
Old World
The Old World consists of those parts of the world known to classical antiquity and the European Middle Ages. It is used in the context of, and contrast with, the "New World" ....
, with only three species known from the Neotropics. Some members of Tessaratomidae exhibit maternal care of eggs and offspring. The defensive chemicals of certain species can cause significant damage if they come into contact with human skin; they may also cause temporary blindness.
All species are exclusively plant-eaters, some of major economic importance as agricultural pests. A few species are also consumed as human food in some countries.
Description
Larger species of Tessaratomidae are known informally as giant shield bugs, giant stink bugs, or inflated stink bugs, but they generally do not have a collective common nameCommon name
A common name of a taxon or organism is a name in general use within a community; it is often contrasted with the scientific name for the same organism...
and are referred to mostly as tessaratomids.
Tessaratomids are ovate to elongate-ovate bugs. They range in size from the smallest members of the tribe
Tribe (biology)
In biology, a tribe is a taxonomic rank between family and genus. It is sometimes subdivided into subtribes.Some examples include the tribes: Canini, Acalypheae, Hominini, Bombini, and Antidesmeae.-See also:* Biological classification* Rank...
Sepinini at 6 to 7 mm (0.236220472440945 to 0.275590551181102 in), to the large Amissus atlas of tribe Tessaratomini at 43 to 45 mm (1.7 to 1.8 in). They are generally quite large and usually exceed 15 mm (0.590551181102362 in) in length.
The head of tessaratomids is generally small and triangular, with the antennae
Antenna (biology)
Antennae in biology have historically been paired appendages used for sensing in arthropods. More recently, the term has also been applied to cilium structures present in most cell types of eukaryotes....
having 4 to 5 segments (though some of them, for example Siphnus, have relatively large heads). The scutellum (Latin for 'little shield', the hard extension of the thorax
Thorax (insect anatomy)
The thorax is the mid section of the insect body. It holds the head, legs, wings and abdomen. It is also called mesosoma in other arthropods....
covering the abdomen
Abdomen
In vertebrates such as mammals the abdomen constitutes the part of the body between the thorax and pelvis. The region enclosed by the abdomen is termed the abdominal cavity...
in hemiptera
Hemiptera
Hemiptera is an order of insects most often known as the true bugs , comprising around 50,000–80,000 species of cicadas, aphids, planthoppers, leafhoppers, shield bugs, and others...
ns) is triangular and does not cover the leathery middle section of the forewing but is often partially covered by the prothorax
Prothorax
The prothorax is the foremost of the three segments in the thorax of an insect, and bears the first pair of legs. Its principal sclerites are the pronotum , the prosternum , and the propleuron on each side. The prothorax never bears wings in extant insects, though some fossil groups possessed...
. The tarsi (the final segments of the legs) have 2 to 3 segments.
Like all hemiptera
Hemiptera
Hemiptera is an order of insects most often known as the true bugs , comprising around 50,000–80,000 species of cicadas, aphids, planthoppers, leafhoppers, shield bugs, and others...
ns, instead of mandibles for chewing, tesseratomids possess a piercing-sucking mouthpart for feeding (known as the rostrum). In tesseratomids, the rostrum has 4 segments.
Tessaratomids are oftentimes vividly colored.
Ecology
All tessaratomids are phytophagous. They generally feed upon plants belonging to the plant orders RosalesRosales
Rosales is an order of flowering plants. It is one of the four orders in the nitrogen fixing clade of the fabids and is sister to a clade consisting of Fagales and Cucurbitales. It contains about 7700 species, distributed into about 260 genera. Rosales comprises nine families, the type family...
and Sapindales
Sapindales
Sapindales is a botanical name for an order of flowering plants. Well-known members of Sapindales include citrus; maples, horse-chestnuts, lychees and rambutans; mangos and cashews; frankincense and myrrh; mahogany and neem....
, and spend most of their lives in tree leaves and stems. They exhibit incomplete metamorphosis and have lifespans that can be several years.
Some tessaratomids guard their eggs and nymphs from predators which may include parasitoid
Parasitoid
A parasitoid is an organism that spends a significant portion of its life history attached to or within a single host organism in a relationship that is in essence parasitic; unlike a true parasite, however, it ultimately sterilises or kills, and sometimes consumes, the host...
wasp
Wasp
The term wasp is typically defined as any insect of the order Hymenoptera and suborder Apocrita that is neither a bee nor an ant. Almost every pest insect species has at least one wasp species that preys upon it or parasitizes it, making wasps critically important in natural control of their...
s and assassin bugs.
Life cycle
The eggs of tessaratomids are barrel-shaped or globular. The eggs exhibit a ring of small protuberances, known as micropylar process, which permit entry of sperm for fertilization into the eggs (through micropylar canals). They also provide openings for air for the developing embryos.The eggs are laid in compact clusters glued to the leaves of a variety of plants. The laying arrangement can follow a pattern. For example, in Pygoplatys tenangau, the egg clusters are distinctively hexagonal; while in Piezosternum subulatum, they are arranged in two neat rows. The eggs are usually initially white, cream, or yellow in color but can change as the embryo
Embryo
An embryo is a multicellular diploid eukaryote in its earliest stage of development, from the time of first cell division until birth, hatching, or germination...
s inside mature.
Nymphs emerge from the eggs through peristaltic movement
Peristalsis
Peristalsis is a radially symmetrical contraction and relaxation of muscles which propagates in a wave down the muscular tube, in an anterograde fashion. In humans, peristalsis is found in the contraction of smooth muscles to propel contents through the digestive tract. Earthworms use a similar...
s and with the help of an internal nearly H-shaped structure in the egg known as the 'egg burster'.
As in other hemiptera
Hemiptera
Hemiptera is an order of insects most often known as the true bugs , comprising around 50,000–80,000 species of cicadas, aphids, planthoppers, leafhoppers, shield bugs, and others...
ns, tessaratomids are hemimetabolic
Hemimetabolism
Hemimetabolism or hemimetaboly, also called incomplete metamorphosis, is a term used to describe the mode of development of certain insects that includes three distinct stages: the egg, nymph, and the adult stage, or imago. These groups go through gradual changes; there is no pupal stage...
, undergoing incomplete metamorphosis
Metamorphosis
Metamorphosis is a biological process by which an animal physically develops after birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation...
. This means that they do not possess larva
Larva
A larva 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...
l and pupa
Pupa
A pupa is the life stage of some insects undergoing transformation. The pupal stage is found only in holometabolous insects, those that undergo a complete metamorphosis, going through four life stages; embryo, larva, pupa and imago...
l stages. Instead, juvenile tessaratomids (called nymphs
Nymph (biology)
In biology, a nymph is the immature form of some invertebrates, particularly insects, which undergoes gradual metamorphosis before reaching its adult stage. Unlike a typical larva, a nymph's overall form already resembles that of the adult. In addition, while a nymph moults it never enters a...
), hatch directly from the eggs. The nymphs resemble fully grown adults except for size and the absence of wings.
Nymphs usually undergo four to five successive stages of moltings (ecdysis
Ecdysis
Ecdysis is the moulting of the cuticula in many invertebrates. This process of moulting is the defining feature of the clade Ecdysozoa, comprising the arthropods, nematodes, velvet worms, horsehair worms, rotifers, tardigrades and Cephalorhyncha...
), increasing in size and becoming more adult-like with each stage until the final molting. The stages are individually known as instar
Instar
An instar is a developmental stage of arthropods, such as insects, between each molt , until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or assume a new form. Differences between instars can often be seen in altered body proportions, colors, patterns, or...
s, with the earliest stage (just after hatching) being known as the first nymphal instar. Nymphs may also differ significantly from adults in colors and patterns exhibited. In some species, nymphs often exhibit strikingly vibrant colors in contrast to the relative drabness of adults. The colors can also vary between instars.
Mating between adults can last for several hours, with the male and female attached end-to-end.
Maternal care
Maternal careParental investment
In evolutionary biology, parental investment is any parental expenditure that benefits one offspring at a cost to parents' ability to invest in other components of fitness...
is a well-documented presocial behavior
Presociality
Presociality is a phenomenon in which animals exhibit more than just sexual interactions with members of the same species, but fall short of qualifying as eusocial...
among tessaratomids. Egg guarding by adult females were first observed in 1991 by S. Tachikawa among Japanese species of the genera Pygoplatys (subfamily Tessaratominae) and Erga
Erga
Erga Edizioni, is a publishing company situated in Genova . It was founded on 1964 by Marcello Merli.-History:Erga started publishing some books about history and culture of Genova...
(subfamily Oncomerinae).
In 1998, Gogala et al. described tessaratomines of the genus Pygoplatys from Thailand
Thailand
Thailand , officially the Kingdom of Thailand , formerly known as Siam , is a country located at the centre of the Indochina peninsula and Southeast Asia. It is bordered to the north by Burma and Laos, to the east by Laos and Cambodia, to the south by the Gulf of Thailand and Malaysia, and to the...
and Malaysia showing egg guarding behavior. In addition, they were also observed to exhibit another remarkable maternal behavior. A dense cluster of small nymphs were photographed being carried around by adult females. The nymphs were firmly clutching unto the bottom side of the abdomens of the adults and to each other, forming a compact mass. The females seem unimpeded by their burden and were able to walk around normally and even fly. The nymphs, however, were not observed feeding. This behavior is known as "nymphal phoresy" (used adjectivally
Adjective
In grammar, an adjective is a 'describing' word; the main syntactic role of which is to qualify a noun or noun phrase, giving more information about the object signified....
as "phoretic").
In the Indonesia
Indonesia
Indonesia , officially the Republic of Indonesia , is a country in Southeast Asia and Oceania. Indonesia is an archipelago comprising approximately 13,000 islands. It has 33 provinces with over 238 million people, and is the world's fourth most populous country. Indonesia is a republic, with an...
n species Pygoplatys tenangau, females will cover the clutch of 70 to 120 eggs with their bodies after laying them, literally "standing guard" over them. When approached, they will spray defensive liquid at perceived attackers and may buzz their wings. They will not willingly abandon the eggs they are guarding, however, and if picked up will try to hold unto the leaf where their eggs are attached. It usually takes slightly more than two weeks for the eggs to hatch. The hatching process will take up 3 to 4 days, during which the newly hatched nymphs will immediately clamber unto their mother's abdomen. They were observed to remain phoretic for at least 17 days (Magnien et al., 2008).
In the subfamily Oncomerinae, a predominantly Australia
Australia
Australia , officially the Commonwealth of Australia, is a country in the Southern Hemisphere comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands in the Indian and Pacific Oceans. It is the world's sixth-largest country by total area...
n group of large colorful bugs, brooding behavior varies from species which do not practice it at all (exhibited by Musgraveia sulciventris) to adult females carrying first and second instar
Instar
An instar is a developmental stage of arthropods, such as insects, between each molt , until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or assume a new form. Differences between instars can often be seen in altered body proportions, colors, patterns, or...
nymphs on their abdomens (Monteith, 2011).
Adult female oncomerines of the genus Lyramorpha will guard nymphs at least to the second instar.
Oncomerines of the genera Cumare, Garceus, and Peltocopta exhibit the most advanced form of maternal care. Like the previously described Southeast Asian Pygoplatys individuals, the females actually carry young nymphs around on their abdomens. As the nymphs grow older, they eventually separate from their mothers, lose their bright colors, and become more solitary prior to molting into adults. Species which exhibit this behavior often have significantly flattened and expanded abdomens.
Of the Australian oncomerines, the bronze orange bug (Musgraveia sulciventris) is the only species unequivocally documented to lack maternal brooding behavior. This peculiarity might be connected to the unreliability of the food plant availability for the species (Monteith, 2011). Unlike other oncomerines who can only lay one egg clutch for the certain amount of time it takes to care for them, M. sulciventris can produce multiple egg clutches rapidly because females don't have to care for them. This allows M. sulciventris to rapidly expand their population when conditions are favorable.
Defenses
Tesseratomids, like most heteropterans use chemical defenses (allomoneAllomone
An allomone is any chemical substance produced and released by an individual of one species that affects the behaviour of a member of another species to the benefit of the originator but not the receiver...
s), the source of the common name for pentatomoids - 'stink bugs'. When threatened, tessaratomids may squirt a strong jet of caustic liquid up to a distance of 15 to 27 cm (5.9 to 10.6 in).
The chemicals produced by heteroptera
Heteroptera
Heteroptera is a group of about 40,000 species of insects in the Hemiptera. Sometimes called "true bugs", that name more commonly refers to Hemiptera as a whole, and "typical bugs" might be used as a more unequivocal alternative since among the Hemiptera the heteropterans are most consistently and...
ns are usually alkane
Alkane
Alkanes are chemical compounds that consist only of hydrogen and carbon atoms and are bonded exclusively by single bonds without any cycles...
s and aldehyde
Aldehyde
An aldehyde is an organic compound containing a formyl group. This functional group, with the structure R-CHO, consists of a carbonyl center bonded to hydrogen and an R group....
s from gland
Gland
A gland is an organ in an animal's body that synthesizes a substance for release of substances such as hormones or breast milk, often into the bloodstream or into cavities inside the body or its outer surface .- Types :...
s in the thorax
Thorax (insect anatomy)
The thorax is the mid section of the insect body. It holds the head, legs, wings and abdomen. It is also called mesosoma in other arthropods....
. Compounds that are primarily for protection against fellow arthropod
Arthropod
An arthropod is an invertebrate animal having an exoskeleton , a segmented body, and jointed appendages. Arthropods are members of the phylum Arthropoda , and include the insects, arachnids, crustaceans, and others...
s (to which they are lethal). However, the defensive chemicals of tessaratomids (particularly that of Tessaratoma papillosa and Musgraveia sulciventris) are notable for being one of the most debilitating to vertebrate
Vertebrate
Vertebrates are animals that are members of the subphylum Vertebrata . Vertebrates are the largest group of chordates, with currently about 58,000 species described. Vertebrates include the jawless fishes, bony fishes, sharks and rays, amphibians, reptiles, mammals, and birds...
s, probably a defense specifically aimed against birds. They can cause damage to human skin and even cause temporary blindness if sprayed unto the eyes.
In Lyramoprha parens, nymphs are also known to be highly gregarious, forming massed feeding groups and moving to new feeding sites in close-packed groups. This behavior, along with their bright colorations and stink glands is believed to help in discouraging potential predators.
Aggregation behavior is also common in adults in some species. Aside from combined chemical defenses, other possible benefits of aggregation include better mating opportunities and shelter, greater retention of moisture and heat, and a possible sense of security. Lone bugs in some species were known to be more likely to be skittish than bugs in groups. However, aggregation can also increase the threats of disease
Disease
A disease is an abnormal condition affecting the body of an organism. It is often construed to be a medical condition associated with specific symptoms and signs. It may be caused by external factors, such as infectious disease, or it may be caused by internal dysfunctions, such as autoimmune...
s, parasites, and parasitoid
Parasitoid
A parasitoid is an organism that spends a significant portion of its life history attached to or within a single host organism in a relationship that is in essence parasitic; unlike a true parasite, however, it ultimately sterilises or kills, and sometimes consumes, the host...
s.
If all these defenses fail, tessaratomids will escape predators either by flying away or dropping to the ground (except in cases of females guarding eggs as discussed above).
Natural enemies
Natural enemiesPredation
In ecology, predation describes a biological interaction where a predator feeds on its prey . Predators may or may not kill their prey prior to feeding on them, but the act of predation always results in the death of its prey and the eventual absorption of the prey's tissue through consumption...
of tessaratomids include several tiny parasitoid
Parasitoid
A parasitoid is an organism that spends a significant portion of its life history attached to or within a single host organism in a relationship that is in essence parasitic; unlike a true parasite, however, it ultimately sterilises or kills, and sometimes consumes, the host...
wasp
Wasp
The term wasp is typically defined as any insect of the order Hymenoptera and suborder Apocrita that is neither a bee nor an ant. Almost every pest insect species has at least one wasp species that preys upon it or parasitizes it, making wasps critically important in natural control of their...
s as well as other hemiptera
Hemiptera
Hemiptera is an order of insects most often known as the true bugs , comprising around 50,000–80,000 species of cicadas, aphids, planthoppers, leafhoppers, shield bugs, and others...
ns.
Parasitoid wasps that parasitize tessaratomids usually come from the families Eupelmidae
Eupelmidae
Eupelmidae is a family of parasitic wasps in the superfamily Chalcidoidea. The group is apparently polyphyletic, though the different subfamilies may each be monophyletic, and may be elevated to family status in the near future. As presently defined, there are over 905 described species in 45 genera...
, Scelionidae
Scelionidae
The Hymenopteran family Scelionidae is a very large cosmopolitan group of exclusively parasitoid wasps, mostly small , often black, often highly sculptured, with elbowed antennae that have an 9- or 10-segmented flagellum...
, and Encyrtidae
Encyrtidae
Encyrtidae is a large family of parasitic wasps, with some 3710 described species in some 455 genera . The larvae of the majority are primary parasitoids on Hemiptera, though other hosts are attacked, and details of the life history can be variable Encyrtidae is a large family of parasitic wasps,...
. Adult female parasitoid wasps will search out eggs laid by tessaratomoids. Upon finding some, they will thrust their ovipositor
Ovipositor
The ovipositor is an organ used by some animals for oviposition, i.e., the laying of eggs. It consists of a maximum of three pairs of appendages formed to transmit the egg, to prepare a place for it, and to place it properly...
s into them and lay eggs inside. The eggs of parasitoid wasps hatch and develop inside the tessaratomid eggs, feeding on the tessaratomid embryo and eventually killing it. Infested eggs characteristically turn darker in color as the wasp larva matures. After about a week, one or more adult wasps will then emerge from the now empty egg.
Musgraveia sulciventris is parasitized by the wasps Eupelmus poggioni and Telenomus spp.; Tessaratoma javanica by the wasps Anastatus colemani and Anastatus kashmirensis(?); and the lychee giant stink bug Tessaratoma papillosa by the wasps Ooencyrtus phongi, Anastatus spp. (particularly Anastatus japonicus), and Trissolcus spp. (particularly Trissolcus latisulcus).
In tessaratomids considered to be agricultural pests (like the Musgraveia sulciventris and Tessaratoma papillosa), the wasps that parasitize them are being studied as potential biological control agents
Biological pest control
Biological control of pests in agriculture is a method of controlling pests that relies on predation, parasitism, herbivory, or other natural mechanisms...
. In the Fujian
Fujian
' , formerly romanised as Fukien or Huguing or Foukien, is a province on the southeast coast of mainland China. Fujian is bordered by Zhejiang to the north, Jiangxi to the west, and Guangdong to the south. Taiwan lies to the east, across the Taiwan Strait...
, Guangdong
Guangdong
Guangdong is a province on the South China Sea coast of the People's Republic of China. The province was previously often written with the alternative English name Kwangtung Province...
, and Guangxi
Guangxi
Guangxi, formerly romanized Kwangsi, is a province of southern China along its border with Vietnam. In 1958, it became the Guangxi Zhuang Autonomous Region of the People's Republic of China, a region with special privileges created specifically for the Zhuang people.Guangxi's location, in...
provinces of China, mass-reared Anastatus japonicus are being released to combat Tessaratoma papillosa pests in lychee and longan
Longan
Dimocarpus longan, commonly known as the longan, is a tropical tree native to South and Southeast Asia, in the Indomalaya ecozone known for its edible fruit.-Vernacular names:The fruit is known as longan or longyan in English...
crops. The same measures are also reportedly being done in Thailand.
Musgraveia sulciventris is also preyed upon by the predatory pentatomid
Pentatomidae
Pentatomidae, Greek pente meaning five and tomos meaning section, is a family of insects belonging to order hemiptera that includes some of the stink bugs and shield bugs. Their antennae are 5-segmented, which gives the family its scientific name. Their bodies are usually shield-shaped...
Asopus and assassin bugs (family Reduviidae
Reduviidae
Reduviidae is a large, cosmopolitan family of predatory insects in the suborder Heteroptera...
) of the species Pristhesancus papuensis and Pristhesancus plagipennis.
As food
The edible stink bug Encosternum delegorguei is consumed as human food in ZimbabweZimbabwe
Zimbabwe is a landlocked country located in the southern part of the African continent, between the Zambezi and Limpopo rivers. It is bordered by South Africa to the south, Botswana to the southwest, Zambia and a tip of Namibia to the northwest and Mozambique to the east. Zimbabwe has three...
and among the Venda people
Venda people
The Venda are a Southern African people living mostly near the South African-Zimbabwean border.- History :The Venda were originally from either the Congo or the Great Rift Valley, migrating across the Limpopo river during the Bantu expansion.The Venda of today are descendants of many heterogeneous...
of South Africa
South Africa
The Republic of South Africa is a country in southern Africa. Located at the southern tip of Africa, it is divided into nine provinces, with of coastline on the Atlantic and Indian oceans...
. The insects are light green in color and quite large, averaging at 25 mm (0.984251968503937 in) in length. They are most widely known in South Africa as "thongolifha", though they are also known as "tsonônô". In Zimbabwe, they are known as "harurwa" or "harugwa".
Encosternum delegorguei are collected just before dawn when they are least active and are easier to catch. They are caught carefully, taking care not to kill them. The chemicals released by the bugs can often stain the hands of collectors orange if they collect them barehanded. The bugs which died during collection are carefully separated from live bugs. This is because the chemicals stored in the stink glands are unpalatable - being extremely bitter. As dead bugs can not release the remaining chemicals in their bodies, they are deemed unsuitable for consumption and discarded. The remaining live bugs are placed in a bucket with a small amount of warm water. This is then carefully agitated so as to make them release all their defense chemicals in alarm. This is repeated several more times until their stink glands are drained.
The live bugs with their now empty stink glands are then boiled in water. Further sorting is done afterwards. Dead bugs which died before they could release all their chemicals can be distinguished from the 'clean' bugs by their blackened abdomens after boiling. These are also rejected. The remaining bugs are then dried under the sun.
In cases where the bugs were collected dead, another method is used to remove the chemicals. The bugs are beheaded and carefully squeezed so chemicals in their stink glands flow out the severed neck. The liquids secreted are then wiped off and the bugs boiled and sun-dried like the previous procedure.
After removing the wings, the dried bugs can be eaten as is, fried with a little salt, or cooked with a type of porridge called pap
Pap (food)
Pap , a traditional porridge made from mielie-meal , is a staple food of the Bantu inhabitants of South Africa...
. They are believed to be a good source of protein
Protein
Proteins are biochemical compounds consisting of one or more polypeptides typically folded into a globular or fibrous form, facilitating a biological function. A polypeptide is a single linear polymer chain of amino acids bonded together by peptide bonds between the carboxyl and amino groups of...
.
Diminishing harvests of E. delegorguei has been a cause for concern in recent years. It may be due to the decline in the number of available food plants which are being harvested locally for firewood. Studies are being done in South Africa for ways to ensure sustainable harvests of E. delegorguei, as well as for the possibilities of rearing them in captivity for human consumption.
In Thailand
Thailand
Thailand , officially the Kingdom of Thailand , formerly known as Siam , is a country located at the centre of the Indochina peninsula and Southeast Asia. It is bordered to the north by Burma and Laos, to the east by Laos and Cambodia, to the south by the Gulf of Thailand and Malaysia, and to the...
(where a total of 81 insect species are reportedly eaten), large tessaratomids of the genera Pygoplatys and Tessaratoma (T. papillosa and T. javanica) are eaten.
In Laos
Laos
Laos Lao: ສາທາລະນະລັດ ປະຊາທິປະໄຕ ປະຊາຊົນລາວ Sathalanalat Paxathipatai Paxaxon Lao, officially the Lao People's Democratic Republic, is a landlocked country in Southeast Asia, bordered by Burma and China to the northwest, Vietnam to the east, Cambodia to the south and Thailand to the west...
, Tessaratoma quadrata, locally known as "mien kieng" are also eaten. The same species is also eaten among the Galo people of Northeast India
India
India , officially the Republic of India , is a country in South Asia. It is the seventh-largest country by geographical area, the second-most populous country with over 1.2 billion people, and the most populous democracy in the world...
where they are known as "tari". Only adults are consumed. The wings are removed and the bugs eaten raw or cooked into chutney
Chutney
Chutney is a a condiment used in South Asian cuisine that usually contains a spice and vegetable mix.Chutneys are wet or dry, having a coarse to fine texture. The Anglo-Indian loan word refers to fresh and pickled preparations indiscriminately, with preserves often sweetened. At least several...
.
As agricultural pests
Lychee giant stink bugs, Tessaratoma papillosa, are destructive pests of lychee trees (Litchi chinensis) in ChinaChina
Chinese civilization may refer to:* China for more general discussion of the country.* Chinese culture* Greater China, the transnational community of ethnic Chinese.* History of China* Sinosphere, the area historically affected by Chinese culture...
. They also feed on the closely related Sapindaceae
Sapindaceae
Sapindaceae, also known as the soapberry family, is a family of flowering plants in the order Sapindales. There are about 140-150 genera with 1400-2000 species, including maple, horse chestnut and lychee....
fruit trees like longan
Longan
Dimocarpus longan, commonly known as the longan, is a tropical tree native to South and Southeast Asia, in the Indomalaya ecozone known for its edible fruit.-Vernacular names:The fruit is known as longan or longyan in English...
(Dimocarpus longan) and rambutan
Rambutan
The rambutan is a medium-sized tropical tree in the family Sapindaceae, and the fruit of this tree. It is native to Vietnam, Indonesia, the Philippines, Sri Lanka, Malaysia, and elsewhere in Southeast Asia, although its precise natural distribution is unknown. It is closely related to several...
(Nephelium lappaceum). The closely related Tessaratoma quadrata and Tessaratoma javanica are also minor pests of apple
Apple
The apple is the pomaceous fruit of the apple tree, species Malus domestica in the rose family . It is one of the most widely cultivated tree fruits, and the most widely known of the many members of genus Malus that are used by humans. Apple grow on small, deciduous trees that blossom in the spring...
, pear
Pear
The pear is any of several tree species of genus Pyrus and also the name of the pomaceous fruit of these trees. Several species of pear are valued by humans for their edible fruit, but the fruit of other species is small, hard, and astringent....
, and lychee trees.
| Top left: A first instar Instar An instar is a developmental stage of arthropods, such as insects, between each molt , until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or assume a new form. Differences between instars can often be seen in altered body proportions, colors, patterns, or... nymph Nymph (biology) In biology, a nymph is the immature form of some invertebrates, particularly insects, which undergoes gradual metamorphosis before reaching its adult stage. Unlike a typical larva, a nymph's overall form already resembles that of the adult. In addition, while a nymph moults it never enters a... of the lychee giant shield bug (Tessaratoma papillosa) from China China Chinese civilization may refer to:* China for more general discussion of the country.* Chinese culture* Greater China, the transnational community of ethnic Chinese.* History of China* Sinosphere, the area historically affected by Chinese culture... ; Right: Vividly colored older nymph Nymph (biology) In biology, a nymph is the immature form of some invertebrates, particularly insects, which undergoes gradual metamorphosis before reaching its adult stage. Unlike a typical larva, a nymph's overall form already resembles that of the adult. In addition, while a nymph moults it never enters a... of the same species; Bottom left: The drably colored adult. (Photo by Charles Lam) |
||
Bronze orange bugs (Musgraveia sulciventris) are serious pests to citrus
Citrus
Citrus is a common term and genus of flowering plants in the rue family, Rutaceae. Citrus is believed to have originated in the part of Southeast Asia bordered by Northeastern India, Myanmar and the Yunnan province of China...
crops in Australia
Australia
Australia , officially the Commonwealth of Australia, is a country in the Southern Hemisphere comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands in the Indian and Pacific Oceans. It is the world's sixth-largest country by total area...
. They are very large bugs, around 20 mm (0.78740157480315 in) in length, whose native host plants are members of the rue family, Rutaceae
Rutaceae
Rutaceae, commonly known as the rue or citrus family, is a family of flowering plants, usually placed in the order Sapindales.Species of the family generally have flowers that divide into four or five parts, usually with strong scents...
.
In Indonesia
Indonesia
Indonesia , officially the Republic of Indonesia , is a country in Southeast Asia and Oceania. Indonesia is an archipelago comprising approximately 13,000 islands. It has 33 provinces with over 238 million people, and is the world's fourth most populous country. Indonesia is a republic, with an...
, the tessaratomid Pygoplatys tenangau, locally known as the "tenangau", is considered to be one of the most important pests of Damar gardens. Damar gardens are cultivated forests of trees of the genera Shorea
Shorea
Shorea is a genus of about 196 species of mainly rainforest trees in the family Dipterocarpaceae. The genus is named after Sir John Shore, the Governor-General of the British East India Company, 1793-1798. They are native to southeast Asia, from Northern India to Malaysia, Indonesia and the...
, Balanocarpus, or Hopea
Hopea
Hopea is a genus of plant in family Dipterocarpaceae. The genus was named after John Hope, 1725-1786, the first Regius Keeper of the Royal Botanic Garden, Edinburgh. It contains about 104 species, distributed from Sri Lanka and southern India to southern China, and southward throughout Malesia to...
used as a source of Damar resin
Dammar gum
Dammar gum is obtained from the Dipterocarpaceae family of trees in India and East Asia, principally those of the genera Shorea, Balanocarpus or Hopea. Most is produced by tapping trees; however, some is collected in fossilized form from the ground. The gum varies in colour from clear to pale...
. P. tenangau is the only known tessaratomid which feeds on Dipterocarpaceae
Dipterocarpaceae
Dipterocarpaceae is a family of 17 genera and approximately 500 species of mainly tropical lowland rainforest trees. The family name, from the type genus Dipterocarpus, is derived from Greek and refers to the two-winged fruit...
.
In Papua New Guinea
Papua New Guinea
Papua New Guinea , officially the Independent State of Papua New Guinea, is a country in Oceania, occupying the eastern half of the island of New Guinea and numerous offshore islands...
, Agapophyta viridula and Agapophyta similis are regarded as pests of Tephrosia
Tephrosia
Tephrosia is a genus of flowering plants in the pea family, Fabaceae. The generic name is derived from the Greek word τεφρος , meaning "ash-colored," referring to the greyish tint given to the leaves by their dense trichomes.-Uses:...
spp. and pigeon pea
Pigeon pea
The pigeon pea , also known as tropical green pea, toor dāl or arhar dāl , ತೊಗರಿ ಬೇಳೆ kadios , or Congo pea or gungo pea , pois Congo , gandul , gunga pea, or no-eye pea, [Cajanus...
s (Cajanus cajan). Agapophyta bipunctata are known minor pests of coconut
Coconut
The coconut palm, Cocos nucifera, is a member of the family Arecaceae . It is the only accepted species in the genus Cocos. The term coconut can refer to the entire coconut palm, the seed, or the fruit, which is not a botanical nut. The spelling cocoanut is an old-fashioned form of the word...
s (Cocos nucifera) and sago
Sago
Sago is a starch extracted in the spongy center or pith, of various tropical palm stems, Metroxylon sagu. It is a major staple food for the lowland peoples of New Guinea and the Moluccas, where it is called saksak and sagu. A type of flour, called sago flour, is made from sago. The largest supply...
(Metroxylon sagu) as well.
Classification and distribution
Tessaratomidae was first described as a family group by the Swedish entomologist Carl StålCarl Stål
Carl Stål was a Swedish entomologist specialising in Hemiptera.He was born at Karlberg Castle, Stockholm on 21 March 1833 and died at Frösundavik near Stockholm on 13 June 1878. He matriculated at Uppsala University in 1853, studying medicine and passing the medico-philosophical examination in 1857...
in 1864. In 1900, the Hungarian entomologist Géza Horváth
Géza Horváth
Géza Horváth was a Hungarian doctor and entomologist internationally recognized for his work on bugs .He also contributed extensively to the study of Hungarian scale insect fauna...
divided the family into 9 tribes
Tribe (biology)
In biology, a tribe is a taxonomic rank between family and genus. It is sometimes subdivided into subtribes.Some examples include the tribes: Canini, Acalypheae, Hominini, Bombini, and Antidesmeae.-See also:* Biological classification* Rank...
and established a key to determining genera. The English entomologist George Willis Kirkaldy
George Willis Kirkaldy
George Willis Kirkaldy was an English, entomologist who specialised on Hemiptera. He was a Fellow of the Royal Entomological Society....
increased the number of subfamilies under Tessaratomidae to 11 in 1909. Dennis Leston (1955) followed Kirkaldy's system but reclassified some tribes to subtribes. The current classification is based on the work of Pramod Kumar in 1969 who reduced the number of subfamilies to three - Natalicolinae, Oncomerinae, and Tessaratominae; with Tessaratominae being further divided into three tribes - Prionogastrini, Sepinini and Tessaratomini. Subsequent revisions by Sinclair (1989), Rolston et al. (1993), Schuch & Slater (1995), Sinclair (2000), Cassis & Gross (2002), and Rider (2006), are all based upon Kumar's system.
Tessaratomidae is classified under order
Order (biology)
In scientific classification used in biology, the order is# a taxonomic rank used in the classification of organisms. Other well-known ranks are life, domain, kingdom, phylum, class, family, genus, and species, with order fitting in between class and family...
Hemiptera
Hemiptera
Hemiptera is an order of insects most often known as the true bugs , comprising around 50,000–80,000 species of cicadas, aphids, planthoppers, leafhoppers, shield bugs, and others...
(true bugs), suborder Heteroptera
Heteroptera
Heteroptera is a group of about 40,000 species of insects in the Hemiptera. Sometimes called "true bugs", that name more commonly refers to Hemiptera as a whole, and "typical bugs" might be used as a more unequivocal alternative since among the Hemiptera the heteropterans are most consistently and...
, infraorder Pentatomomorpha
Pentatomomorpha
Pentatomomorpha is an infraorder of insects in the true bug order . It unites such animals as the plant bugs , stink bugs , flat bugs , seed bugs , etc. They are closely related to the Cimicomorpha.-Systematics:5 superfamilies are usually placed in the Pentatomomorpha...
, and superfamily
Superfamily
The term superfamily is used to describe several different concepts in different scientific fields:* Superfamily and Superfamily : a level of biological classification; redirects to Taxonomic rank...
Pentatomoidea (shield bugs and stink bugs). It is currently divided into three subfamilies: Natalicolinae (with 8 genera), Oncomerinae (with 15 genera), and Tessaratominae (with 33 genera and one of uncertain placement).
They are mostly found in tropical Africa, Asia, and Oceania though a few species can be found in the Neotropics and Australia
Australia
Australia , officially the Commonwealth of Australia, is a country in the Southern Hemisphere comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands in the Indian and Pacific Oceans. It is the world's sixth-largest country by total area...
. There are about 240 species known.
Listed below are the three subfamilies; their authors and type genera; the tribes, subtribes, and genera classified under them; and their distribution ranges
Range (biology)
In biology, the range or distribution of a species is the geographical area within which that species can be found. Within that range, dispersion is variation in local density.The term is often qualified:...
:
- FamilyFamilyIn human context, a family is a group of people affiliated by consanguinity, affinity, or co-residence. In most societies it is the principal institution for the socialization of children...
Tessaratomidae StålCarl StålCarl Stål was a Swedish entomologist specialising in Hemiptera.He was born at Karlberg Castle, Stockholm on 21 March 1833 and died at Frösundavik near Stockholm on 13 June 1878. He matriculated at Uppsala University in 1853, studying medicine and passing the medico-philosophical examination in 1857...
, 1864 - type genusType genusIn biological classification, a type genus is a representative genus, as with regard to a biological family. The term and concept is used much more often and much more formally in zoology than it is in botany, and the definition is dependent on the nomenclatural Code that applies:* In zoological...
: Tessaratoma Lepeletier & Serville, 1825 - Subfamily Natalicolinae StålCarl StålCarl Stål was a Swedish entomologist specialising in Hemiptera.He was born at Karlberg Castle, Stockholm on 21 March 1833 and died at Frösundavik near Stockholm on 13 June 1878. He matriculated at Uppsala University in 1853, studying medicine and passing the medico-philosophical examination in 1857...
, 1870 - type genus: Natalicolina Spinola, 1850
-
-
-
-
- Cyclogastridea - Equatorial and West Africa
- Elizabetha - Equatorial Africa
- Empysarus - Southern India and Sri Lanka
- Encosternum - Southern Africa
- Haplosterna - Equatorial Africa
- Natalicola - Africa
- Selenymenum - Equatorial and West Africa
- Stevesonius - Central Africa
-
-
-
- Subfamily Oncomerinae StålCarl StålCarl Stål was a Swedish entomologist specialising in Hemiptera.He was born at Karlberg Castle, Stockholm on 21 March 1833 and died at Frösundavik near Stockholm on 13 June 1878. He matriculated at Uppsala University in 1853, studying medicine and passing the medico-philosophical examination in 1857...
, 1870 - type genus: Oncomeris Laporte, 1832
-
-
-
-
- Agapophyta - Australia, Moluccas, New Guinea, Solomon Islands
- Cumare - Australia (Queensland)
- ErgaErgaErga Edizioni, is a publishing company situated in Genova . It was founded on 1964 by Marcello Merli.-History:Erga started publishing some books about history and culture of Genova...
- Australia - Garceus - Australia (Queensland)
- Lyramorpha - Australia, Moluccas, and New Guinea
- Musgraveia - Australia
- Neosalica - Myanmar, China, India, Sumatra, and Vietnam
- Oncomeris - Australia, Lesser Sunda Islands, Moluccas, New Guinea, Sulawesi
- Peltocopta - Australia
- Piezosternum - Africa, Cape Verde Islands, Central America and the Caribbean, Madagascar, South America
- Plisthenes - Australia, New Guinea, Solomon Islands, Southeast Asia
- Rhoecus - Australia
- Stilida - Australia
- Tamolia - New Guinea
- Tibiospina - Australia (Queensland)
-
-
-
- Subfamily Tessaratominae StålCarl StålCarl Stål was a Swedish entomologist specialising in Hemiptera.He was born at Karlberg Castle, Stockholm on 21 March 1833 and died at Frösundavik near Stockholm on 13 June 1878. He matriculated at Uppsala University in 1853, studying medicine and passing the medico-philosophical examination in 1857...
, 1865 - type genus: Tessaratoma Lepeletier & Serville, 1825
-
-
- TribeTribe (biology)In biology, a tribe is a taxonomic rank between family and genus. It is sometimes subdivided into subtribes.Some examples include the tribes: Canini, Acalypheae, Hominini, Bombini, and Antidesmeae.-See also:* Biological classification* Rank...
Prionogastrini StålCarl StålCarl Stål was a Swedish entomologist specialising in Hemiptera.He was born at Karlberg Castle, Stockholm on 21 March 1833 and died at Frösundavik near Stockholm on 13 June 1878. He matriculated at Uppsala University in 1853, studying medicine and passing the medico-philosophical examination in 1857...
, 1870 - type genus: Prionogaster StålCarl StålCarl Stål was a Swedish entomologist specialising in Hemiptera.He was born at Karlberg Castle, Stockholm on 21 March 1833 and died at Frösundavik near Stockholm on 13 June 1878. He matriculated at Uppsala University in 1853, studying medicine and passing the medico-philosophical examination in 1857...
, 1853
-
-
- Prionogaster - South Africa
-
- Tribe Sepinini Horváth, 1900 - type genusType genusIn biological classification, a type genus is a representative genus, as with regard to a biological family. The term and concept is used much more often and much more formally in zoology than it is in botany, and the definition is dependent on the nomenclatural Code that applies:* In zoological...
: Sepina Signoret, 1861
-
- Subtribe Platytataria Horváth, 1900 - type genus: Platytatus Bergroth, 1892
-
- Platytatus - Madagascar
- Subtribe Sepinaria Horváth, 1900 - type genus: Sepina Signoret, 1861
-
- Ipamu - Central Africa
- Malgassus - Madagascar
- Pseudosepina - Madagascar
- Rhynchotmetus - Madagascar
- Sepina - Madagascar, Seychelles
- Tribe Tessaratomini StålCarl StålCarl Stål was a Swedish entomologist specialising in Hemiptera.He was born at Karlberg Castle, Stockholm on 21 March 1833 and died at Frösundavik near Stockholm on 13 June 1878. He matriculated at Uppsala University in 1853, studying medicine and passing the medico-philosophical examination in 1857...
, 1864 - type genus: Tessaratoma Lepeletier & Serville, 1825
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- Subtribe Eusthenaria StålCarl StålCarl Stål was a Swedish entomologist specialising in Hemiptera.He was born at Karlberg Castle, Stockholm on 21 March 1833 and died at Frösundavik near Stockholm on 13 June 1878. He matriculated at Uppsala University in 1853, studying medicine and passing the medico-philosophical examination in 1857...
, 1870 - type genus: Eusthenes Laporte, 1832
- Subtribe Eusthenaria Stål
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- Anacanthopus - Philippines
- Asiarcha - China, India, Indochina
- Aurungabada - India (Bombay)
- Candace - West Africa
- Carpona - China, India, Southeast Asia
- Dalcantha - India, Southeast Asia
- Eurostus - East Asia, South Asia, Southeast Asia
- Eurypleura - Indonesia (Java and Sumatra)
- Eusthenes - East Asia, South Asia, Southeast Asia
- Eusthenimorpha - China
- Mattiphus - China, Indochina, Philippines, Sri Lanka, Sulawesi, Sumatra
- Megaedoeum - West Africa
- Origanaus - China
- Pseudopycanum - Malaysia
- Pycanum - East Asia, South Asia, Southeast Asia
- Sanganus - Borneo, New Guinea, Sumatra
- Serrocarpona - Sulawesi
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- Subtribe Tessaratomaria StålCarl StålCarl Stål was a Swedish entomologist specialising in Hemiptera.He was born at Karlberg Castle, Stockholm on 21 March 1833 and died at Frösundavik near Stockholm on 13 June 1878. He matriculated at Uppsala University in 1853, studying medicine and passing the medico-philosophical examination in 1857...
, 1864 - type genus: Tessaratoma Lepeletier & Serville, 1825
- Subtribe Tessaratomaria Stål
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- Acidosterna - Malaysia, Sumatra
- Amissus - Southeast Asia
- Embolosterna - East Asia, South Asia, Southeast Asia
- Enada - Borneo
- Hypencha - Southeast Asia
- Mucanum - Southeast Asia
- Pygoplatys - South and Southeast Asia
- Siphnus - Southeast Asia
- Tessaratoma - Africa, Australia, South Asia, East Asia, Southeast Asia
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- Tribe Notopomini Horváth, 1900 incertae sedisIncertae sedis, is a term used to define a taxonomic group where its broader relationships are unknown or undefined. Uncertainty at specific taxonomic levels is attributed by , , and similar terms.-Examples:*The fossil plant Paradinandra suecica could not be assigned to any...
- type genus: Notopomis Montandon, 1894
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- Notopomus - Malaysia (Pinang Island)
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- Tribe Notopomini Horváth, 1900 incertae sedis
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Fossil record
A fossil specimen, named Tesseratomoides maximus and thought to belong to Tessaratomidae, was recovered in 1967 from the EoceneEocene
The Eocene Epoch, lasting from about 56 to 34 million years ago , is a major division of the geologic timescale and the second epoch of the Paleogene Period in the Cenozoic Era. The Eocene spans the time from the end of the Palaeocene Epoch to the beginning of the Oligocene Epoch. The start of the...
of Germany
Germany
Germany , officially the Federal Republic of Germany , is a federal parliamentary republic in Europe. The country consists of 16 states while the capital and largest city is Berlin. Germany covers an area of 357,021 km2 and has a largely temperate seasonal climate...
; but the specimen was published with no formal description and is thus unacceptable as a valid taxon
Taxon
|thumb|270px|[[African elephants]] form a widely-accepted taxon, the [[genus]] LoxodontaA taxon is a group of organisms, which a taxonomist adjudges to be a unit. Usually a taxon is given a name and a rank, although neither is a requirement...
. Another fossil genus, Latahcoris, from the Miocene
Miocene
The Miocene is a geological epoch of the Neogene Period and extends from about . The Miocene was named by Sir Charles Lyell. Its name comes from the Greek words and and means "less recent" because it has 18% fewer modern sea invertebrates than the Pliocene. The Miocene follows the Oligocene...
Latah Formation
Latah Formation
The Latah Formation is a series of late Miocene lacustrine sedimentary deposits which outcrop in eastern Washington and northwestern Idaho. Its lake beds are interbedded with igneous rock of the Columbia River Basalt Group. The formation was originally named from a site in Spokane, Washington by...
of Idaho
Idaho
Idaho is a state in the Rocky Mountain area of the United States. The state's largest city and capital is Boise. Residents are called "Idahoans". Idaho was admitted to the Union on July 3, 1890, as the 43rd state....
, was described in 1931 by T.D.A. Cockerell.
Phylogeny
A study on the phylogenetic relationships of the superfamilySuperfamily
The term superfamily is used to describe several different concepts in different scientific fields:* Superfamily and Superfamily : a level of biological classification; redirects to Taxonomic rank...
Pentatomoidea in 2008 hints that Tessaratomidae and Dinidoridae represented a monophyletic group. However, the difficulty in securing enough materials for examination for both groups leaves this as yet unresolved.
Below is the morphological
Morphology (biology)
In biology, morphology is a branch of bioscience dealing with the study of the form and structure of organisms and their specific structural features....
unweighted tree
Phylogenetic tree
A phylogenetic tree or evolutionary tree is a branching diagram or "tree" showing the inferred evolutionary relationships among various biological species or other entities based upon similarities and differences in their physical and/or genetic characteristics...
of the superfamily Pentatomoidea after Grazia et al. (2008). Tessaratomidae is in bold. Both Dinidoridae and Tessaratomidae are shown in dotted lines, signifying uncertain status.
External links
- Illustrated catalog of Tessaratomidae, maintained by Philippe Magnien
- Tessaratomids of Australia from Save Our Waterways Now
- Family Tessaratomidae - Large Stink Bugs from Brisbane Insects and Spiders
- Pentatomoidea Home page, maintained by David A. Rider