Tessaratomidae

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 name 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 , to the large Amissus atlas of tribe Tessaratomini at . They are generally quite large and usually exceed 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, the hard extension of the thorax covering the abdomen, is triangular. It 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. Tessaratomids are most reliably distinguished from pentatomids by having six exposed abdominal spiracles instead of five. Like all hemipterans, instead of mandibles for chewing, tesseratomids possess a piercing-sucking mouthpart for feeding (known as the rostrum). In tesseratomids, the rostrum has 4 segments. Bright greens and reds are common colorations. ==Ecology==

All tessaratomids are phytophagous. They generally feed upon plants belonging to the plant orders Rosales 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 wasps and assassin bugs. Life cycle '') mating. 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. Nymphs emerge from the eggs through peristaltic movements and with the help of an internal nearly H-shaped structure in the egg known as the 'egg burster'. As in other hemipterans, tessaratomids are hemimetabolic, undergoing incomplete metamorphosis. This means that they do not possess larval and pupal 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 instars, with the earliest stage (just after hatching) being known as the first nymphal instar. This behavior is known as "nymphal phoresy" (used adjectivally as "phoretic"). In the Indonesian 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 onto 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 Australian 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. Adult female oncomerines of the genus Lyramorpha will guard nymphs at least to the second instar. This allows M. sulciventris to rapidly expand their population when conditions are favorable. 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 . They can cause damage to human skin and even cause temporary blindness if sprayed unto the eyes. 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 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. ==Economic significance==

As food '') being sorted after they had been boiled and dried. The insects are light green in color and quite large, averaging at 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 die 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. 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 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. As agricultural pests Lychee giant stink bugs, Tessaratoma papillosa, are destructive pests of lychee trees (Litchi chinensis) in China. 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 in length, whose native host plants are members of the rue family, Rutaceae. 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. ==Classification and distribution==

Tessaratomidae was first described as a family group by the Swedish entomologist Carl Stål in 1864. 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: ===Natalicolinae=== Authority: Stål, 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 ===Oncomerinae=== Authority: Stål, 1870 - type genus: Oncomeris Laporte, 1832 • Agapophyta - Australia, Moluccas, New Guinea, Solomon Islands • Cumare - Australia (Queensland) • Erga - 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 • Sciadiocoris - Papua New Guinea • Stilida - Australia • Tamolia - New Guinea • Tibiospina - Australia (Queensland) ===Tessaratominae=== Authority: Stål, 1865 - type genus: Tessaratoma Lepeletier & Serville, 1825 ;Tribe Prionogastrini Stål, 1870 - type genus: Prionogaster Stål, 1853 • Prionogaster - South Africa ;Tribe Sepinini Horváth, 1900 - type genus: Sepina Signoret, 1861 • Subtribe Platytatina Horváth, 1900 - type genus: Platytatus Bergroth, 1892 • Platytatus - Madagascar • Subtribe Sepinina Horváth, 1900 - type genus: Sepina Signoret, 1861 • Ipamu - Central Africa • Malgassus - Madagascar • Pseudosepina - Madagascar • Rhynchotmetus - Madagascar • Sepina - Madagascar, Seychelles ;Tribe Tessaratomini Stål, 1864 Selected genera: • Subtribe Eusthenina Stål, 1870 - type genus: Eusthenes Laporte, 1832 • Asiarcha - China, India, Indochina • Candace - Sub-saharan Africa • Eurostus - East Asia, South Asia, Southeast Asia • Eusthenes - East Asia, South Asia, Southeast Asia • Mattiphus - China, Indochina, Philippines, Sri Lanka, Sulawesi, Sumatra • Subtribe Tessaratomina Stål, 1864 - type genus: Tessaratoma Lepeletier & Serville, 1825 • Pygoplatys - South and Southeast Asia • Siphnus - Southeast Asia • Tessaratoma - Africa, Australia, South Asia, East Asia, Southeast Asia ;Tribe Notopomini Horváth, 1900 incertae sedis - type genus: Notopomis Montandon, 1894 • Notopomus - Malaysia (Pinang Island) ==Evolution==

Fossil record A fossil specimen, named Tesseratomoides maximus and thought to belong to Tessaratomidae, was recovered in 1967 from the Eocene of Germany; but the specimen was published with no formal description and is thus unacceptable as a valid taxon. Phylogeny A study on the phylogenetic relationships of the superfamily 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. ==See also==