Tessaratomidae

Tessaratomidae
An adult Pycanum rubens on a leaf.
Pycanum rubens, an adult from the Gunung Mulu National Park, Malaysia
(Photo by Alan Cressler)
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hemiptera
Suborder: Heteroptera
Infraorder: Pentatomomorpha
Superfamily: Pentatomoidea
Family: Tessaratomidae
Stål, 1864
Type genus
Tessaratoma
Lepeletier & Serville, 1825
Subfamilies

See text

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.

Description

Larger species of Tessaratomidae are known informally as giant shield bugs, giant stink bugs, or inflated stink bugs,[1] 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 6 to 7 mm (0.24 to 0.28 in),[2] to the large Amissus atlas of tribe Tessaratomini at 43 to 45 mm (1.7 to 1.8 in).[3] They are generally quite large and usually exceed 15 mm (0.59 in) in length.[4]

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 hemipterans) 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.[5] They 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.[6]

Tessaratomids are oftentimes vividly colored.[7]

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.[6][8] 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

Two bronze orange bugs (Musgraveia sulciventris) 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.[9] The laying arrangement can follow a pattern. For example, in Pygoplatys tenangau, the egg clusters are distinctively hexagonal;[10] while in Piezosternum subulatum, they are arranged in two neat rows.[11] The eggs are usually initially white, cream, or yellow in color but can change as the embryos inside mature.[10][11][12]

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'.[9]

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.[13] 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.[13]

Mating between adults can last for several hours, with the male and female attached end-to-end.[10]

Life cycle of the bronze orange bug (Musgraveia sulciventris)
A cluster of 13 small spherical eggs glued together on a twig. Visible through each are the pair of eyes of the developing embryos, except for an unfertilized egg.
(Photo by Emily Sephton)
Small flattened ovoid bug wandering on a twig. Empty eggs lie below them with holes at the tops.
(Photo by Emily Sephton)
An oval dorsoventrally flattened nymph of the bronze orange bug on a citrus leaf.
(Photo by Andy Burton)
A bronze orange bug clinging to the underside of an orange leaf. The shape of its body is distinctly shield-like.
(Photo by Jan Anderson)
Left: A cluster of bronze orange bug eggs. You can make out the embryos through the clear egg membranes, as well as the small ring of micropylar processes on each egg. The second egg from the bottom right is unfertilized and remains a murky green; Center left: Nymphs emerging from the eggs. Early instars of bronze orange bugs are bright green in color; Center right: A fourth or fifth instar nymph resting on a citrus leaf. It is now brilliantly orange in color with black margins and a small black dot at the center of its body; Right: An adult bronze orange bug on the underside of a citrus leaf. The adults are much drabber in color than the nymphs. Below it is also a green third instar nymph

Maternal care

A distinctly rectangular and dorsoventrally flattened nymph crawling on some mossy twigs.
The brilliantly colored and strangely shaped nymphs of Pycanum rubens bear little resemblance to the adults.
(Photo by Alan Cressler)

Maternal care 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).[14][15]

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.[16] This behavior is known as "nymphal phoresy" (used adjectivally as "phoretic").[10][12][17]

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.[18] 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).[10]

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 (Monteith, 2011).[19]

Adult female oncomerines of the genus Lyramorpha will guard nymphs at least to the second instar.[19]

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.[19] Species which exhibit this behavior often have significantly flattened and expanded abdomens.[4]

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.[15] This allows M. sulciventris to rapidly expand their population when conditions are favorable.[19]

Defenses

Tesseratomids, like most heteropterans use chemical defenses (allomones),[20] the source of the common name for pentatomoids - 'stink bugs'.[13] When threatened, tessaratomids may squirt a strong jet of caustic liquid up to a distance of 15 to 27 centimetres (5.9 to 10.6 in).[21]

The chemicals produced by heteropterans are usually alkanes and aldehydes from glands in the thorax. Compounds that are primarily for protection against fellow arthropods (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 vertebrates, probably a defense specifically aimed against birds.[21] They can cause damage to human skin and even cause temporary blindness if sprayed unto the eyes.[10]

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.[19]

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 diseases, parasites, and parasitoids.[22]

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).[10][12]

Natural enemies

An assassin bug walking on a fine metal mesh.
Two small wasps rest on the nearby plant stalks while several more are emerging from the host eggs.
Left: The Australian common assassin bug Pristhesancus plagipennis is a predator of the bronze orange bug Musgraveia sulciventris.
Right: Parasitoid wasps of the genus Trissolcus emerging from the eggs of a pentatomid, Graphosoma lineatum.

Natural enemies of tessaratomids include several tiny parasitoid wasps as well as other hemipterans.[23]

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 ovipositors into them and lay eggs inside.[24] 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.[25]

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).[23]

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.[26][27] 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.[12]

Musgraveia sulciventris is also preyed upon by the predatory pentatomid Asopus and assassin bugs (family Reduviidae) of the species Pristhesancus papuensis and Pristhesancus plagipennis.[23]

Economic significance

As food

A woman wearing protective gloves and holding a small pail reaches out to the branches of a small tree to catch stink bugs.
Hundreds of live stink bugs inside a bucket.
One woman stirring the stink bugs in the bucket carefully as another pours a small amount of water into it.
Edible stink bugs (Encosternum delegorguei) are harvested during early morning. They are placed in a bucket with a small amount of warm water and agitated to release their unpalatable defensive chemicals.[22]
Dead and dried stink bugs spread out on a nylon grain bag and being sorted
Edible stink bugs (Encosternum delegorguei) being sorted after they had been boiled and dried.[22]

The edible stink bug Encosternum delegorguei is consumed as human food in Zimbabwe and among the Venda people of South Africa.[22][28] The insects are light green in color and quite large, averaging at 25 mm (0.98 in) in length. They are most widely known in South Africa as "thongolifha", though they are also known as "tsonônô".[29] In Zimbabwe, they are known as "harurwa" or "harugwa".[30]

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.[31] 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.[31][32]

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.[31] These are also rejected. The remaining bugs are then dried under the sun.[32]

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.[32]

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.[33]

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.[22]

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.[34]

In Laos, Tessaratoma quadrata, locally known as "mien kieng" are also eaten.[35] 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.[36]

As agricultural pests

Lychee giant stink bugs, Tessaratoma papillosa, are destructive pests of lychee trees (Litchi chinensis) in China.[6][37] 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.[26][38]

Top left: A first instar nymph of the lychee giant shield bug (Tessaratoma papillosa) from China; Right: Vividly colored older nymph of the same species; Bottom left: The drably colored adult.
(Photo by Charles Lam)

Bronze orange bugs (Musgraveia sulciventris) are serious pests to citrus crops in Australia. They are very large bugs, around 20 mm (0.79 in) in length, whose native host plants are members of the rue family, Rutaceae.[5]

In Indonesia, the tessaratomid Pygoplatys tenangau, locally known as the "tenangau", is considered to be one of the most important pests of Damar gardens.[39] 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.[10]

In Papua New Guinea, Agapophyta viridula and Agapophyta similis are regarded as pests of Tephrosia spp. and pigeon peas (Cajanus cajan). Agapophyta bipunctata are known minor pests of coconuts (Cocos nucifera) and sago (Metroxylon sagu) as well.[40]

Classification and distribution

Tessaratomidae was first described as a family group by the Swedish entomologist Carl Stål in 1864.[5] 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.[6][41]

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).[41]

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.[42]

Listed below are the three subfamilies; their authors and type genera; the tribes, subtribes, and genera classified under them; and their distribution ranges:[43]

  • Subfamily Natalicolinae 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[44]
  • Subfamily Oncomerinae 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
  • Stilida - Australia
  • Tamolia - New Guinea
  • Tibiospina - Australia (Queensland)
  • Subfamily Tessaratominae Stål, 1865 - type genus: Tessaratoma Lepeletier & Serville, 1825
  • Prionogaster - South Africa
  • Tribe Sepinini Horváth, 1900 - type genus: 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 Stål, 1864 - type genus: Tessaratoma Lepeletier & Serville, 1825
  • Subtribe Eusthenaria Stål, 1870 - type genus: Eusthenes Laporte, 1832
  • 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
  • Subtribe Tessaratomaria Stål, 1864 - type genus: Tessaratoma Lepeletier & Serville, 1825
  • 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
  • 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.[5] Another fossil genus, Latahcoris, from the Miocene Latah Formation of Idaho, was described in 1931 by T.D.A. Cockerell.[45][46]

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.[17]

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.[47]

      ←      
             

Urostylididae


             
             

Saileriolidae


             
             

Acanthosomatidae


             
             
             

Tessaratomidae


             

Dinidoridae


  Cydnidae sensu lato  
             

Cydnidae


             

Thaumastellidae


             

Parastrachiinae



             
             

Corimelaenidae


             

Lestoniidae


             

Phloeidae


             
             

Scutelleridae


             

Plataspididae



             

Pentatomidae


             

Canopidae


             

Megarididae









See also

References

  1. ↑ Cedric Gillott (1995). Entomology. Springer. p. 604. ISBN 978-0-306-44967-3.
  2. ↑ Philippe Magnien (December 12, 2007). "Sepina aberrans (Schouteden, 1905)". Heteroptera, un site pour les punaises. Retrieved April 30, 2011.
  3. ↑ タイヨロイオオカメムシ (in Japanese). 世界のカメムシ. December 12, 2007. Retrieved April 30, 2011.
  4. ↑ 4.0 4.1 Mike Picker, Charles Griffiths, & Alan Weaving (2004). Field guide to insects of South Africa. Struik. p. 134. ISBN 978-1-77007-061-5.
  5. ↑ 5.0 5.1 5.2 5.3 Randall T. Schuh & James Alexander Slater (1995). True bugs of the world (Hemiptera:Heteroptera): classification and natural history. Cornell University Press. p. 241. ISBN 978-0-8014-2066-5.
  6. ↑ 6.0 6.1 6.2 6.3 Jen-Zon Ho & Young-Fa Chen (2010). "A Review of the Family Tessaratomidae(Hemiptera: Pentatomoidea) of Taiwan with Descriptions of Newly Recorded Two Genera and Five Species (台灣荔蝽科昆蟲二新紀錄屬及五新紀錄種)". Taiwan Journal of Biodiversity (台灣生物多樣性研究) (in English with Chinese translations) 12 (4): 393–406. Retrieved April 30, 2011.
  7. ↑ Augustus Daniel Imms, Owain Westmacott Richards, & Richard Gareth Davies (1977). Imms' General textbook of entomology. Volume 2. Springer. pp. 741–742. ISBN 978-0-412-61390-6.
  8. ↑ G. Cassis & Gordon F. Gross (2002). Zoological catalogue of Australia: Hemiptera: Heteroptera (Pentatomomorpha). Csiro Publishing. p. 604. ISBN 978-0-643-06875-9.
  9. ↑ 9.0 9.1 T.R.E. Southwood (1956). "The Structure of the Eggs of the Terrestrial Heteroptera and its Relationship to the Classification of the Group". The Transactions of the Royal Entomological Society of London (Royal Entomological Society) 108 (6): 163–221. Retrieved May 1, 2011.
  10. ↑ 10.0 10.1 10.2 10.3 10.4 10.5 10.6 10.7 Philippe Magnien, Koen Smets, Dominique Pluot-Sigwalt, and Jérôme Constant (2008). "A new species of Pygoplatys Dallas (Heteroptera, Tessaratomidae) from the Damar agroforests in Sumatra: description, immatures and biology". Nouvelle Revue d'Entomologie (Paris) 24 (2): 99–112. Retrieved April 30, 2011.
  11. ↑ 11.0 11.1 Jonathan Figueroa Jiménez & Nohely Trabal. "Piezosternum subulatum (Thunberg 1783)". Department of Biology, University of Puerto Rico at Mayagüez. Retrieved May 1, 2011.
  12. ↑ 12.0 12.1 12.2 12.3 John L. Capinera (2008). Encyclopedia of entomology. Springer. p. 2749. ISBN 978-1-4020-6242-1.
  13. ↑ 13.0 13.1 13.2 Tony, Sandy & Peter Chew (August 7, 2010). "Family Tessaratomidae - Large Stink Bugs". Brisbane Insects and Spiders. Retrieved May 1, 2011.
  14. ↑ S. Tachikawa (1991). Studies on the Subsociality of Japanese Heteroptera(in Japanese). Tokyo Agricultural Press, Tokyo.
  15. ↑ 15.0 15.1 Jae C. Choe & Bernard J. Crespi (1997). The evolution of social behavior in insects and arachnids. Cambridge University Press. pp. 94, 103–105. ISBN 978-0-521-58977-2.
  16. ↑ Matija Gogala, Hoi-Sen Yong, & Carsten Brühl (1998). "Maternal care in Pygoplatys bugs (Heteroptera: Tessaratomidae)". European Journal of Entomology (Czech Entomological Society) 95: 311–315. ISSN 1210-5759. Retrieved April 30, 2011.
  17. ↑ 17.0 17.1 Jocelia Grazia, Randall T. Schuhb, & Ward C. Wheeler (2008). "Phylogenetic relationships of family groups in Pentatomoidea based on morphology and DNA sequences (Insecta: Heteroptera)". Cladistics (Wiley-Blackwell) 24: 932–976. doi:10.1111/j.1096-0031.2008.00224.x. Retrieved April 27, 2011.
  18. ↑ Kyle R. Conrad (2009). "Parental Care in Arthropods: Bracket Organization and Discussion of Behaviors". p. 4. Retrieved April 30, 2011.
  19. ↑ 19.0 19.1 19.2 19.3 19.4 Geoff B. Monteith (2011). "Maternal Care, Food Plants and Distribution of Australian Oncomerinae (Hemiptera: Heteroptera: Tessaratomidae)". Australian Entomologist 38 (1): 37–48. Retrieved April 30, 2011.
  20. ↑ Ashraf M. El-Sayed (November 6, 2009). "Genus Compounds: Tessaratoma, Family: Tessaratomidae, Order: Heteroptera". The Pherobase. Retrieved May 1, 2011.
  21. ↑ 21.0 21.1 Jan Raška (2009). Function of metathoracic scent glands in terrestrial Heteroptera (Bachelor thesis). Univerzita Karlova v Praze. Retrieved May 1, 2011.
  22. ↑ 22.0 22.1 22.2 22.3 22.4 C. M. Dzerefos, E. T. F. Witkowski, & R. Toms (2009). "Life-history traits of the edible stinkbug, Encosternum delegorguei (Hem., Tessaratomidae), a traditional food in southern Africa". Journal of Applied Entomology 133: 739–759. doi:10.1111/j.1439-0418.2009.01425.x.
  23. ↑ 23.0 23.1 23.2 David A. Rider (January 14, 2008). "Natural Enemies Index". Department of Entomology, North Dakota State University. Retrieved May 1, 2011.
  24. ↑ Liu YuFang & Gu DeXiang (2000). "Host-searching behaviour of the egg parasitoid Anastatus japonicus". Chinese Journal of Biological Control 16 (1): 1–4. ISSN 1000-1034. Retrieved May 1, 2011.
  25. ↑ "Parasitoids". Integrated Pest Management (IPM) Thailand. Retrieved May 5, 2011.
  26. ↑ 26.0 26.1 Vasudha Gautam, Suneel Kumar & M.A. Khan (2004). "Natural egg parasitism of litchi stink bug, Tessaratoma javanica (Thunberg) (Hemiptera: Pentatomidae) in India.". Journal of Biological Control 18 (2): 199–201. ISSN 0971-930X. Retrieved May 1, 2011.
  27. ↑ Liu YuFang, Zhang GuRen, Gu DeXiang (2008). "The Natural Control Effect of Egg Parasitoids on Litchi Stink Bug Tessaratoma papillosa (Heteroptera: Pentatomidae) in LItchi Orchard". Insect Science 7 (4): 322–328. doi:10.1111/j.1744-7917.2000.tb00230.x.
  28. ↑ Cathy Dzerefos, David A. Rider (July 28, 2010). "Thongolifha (Encosternum delegorguei)". Department of Entomology, North Dakota State University. Retrieved April 30, 2011.
  29. ↑ Cathy Dzerefos. "Traditional Cuisine With A Difference". Kruger National Park. Retrieved April 30, 2011.
  30. ↑ Pathisa Nyathi (2005). Zimbabwe's cultural heritage. African Books Collective. p. 94. ISBN 978-0-7974-2897-3.
  31. ↑ 31.0 31.1 31.2 Phillip Chidavaenzi (September 7, 2010). "Harurwa – more than food in Bikita". News Day. Retrieved April 30, 2011.
  32. ↑ 32.0 32.1 32.2 Rob Toms & Mashudu Thagwana (2002). "Eat Your Bugs!". Science in Africa. Retrieved April 30, 2011.
  33. ↑ Leah Snow Teffo (2006). Nutritional and Medicinal value of the edible stinkbug, Encosternum delegorguei Spinola consumed in the Limpopo Province of South Africa and its host plant Dodonaea viscosa Jacq. var. angustifolia (Ph.D. thesis). University of Pretoria. Retrieved April 30, 2011.
  34. ↑ Yupa Hanboonsong (2010). "Edible insects and associated food habits in Thailand". In Patrick B. Durst, Dennis V. Johnson, Robin N. Leslie, & Kenichi Shono. Forest insects as food: humans bite back. Food and Agricultural Organization of the United Nations, Regional Office for Asia and the Pacific. ISBN 978-92-5-106488-7.
  35. ↑ Somkhit Boulidam (2010). "Edible insects in a Lao market economy". In Patrick B. Durst, Dennis V. Johnson, Robin N. Leslie, & Kenichi Shono. Forest insects as food: humans bite back. Food and Agricultural Organization of the United Nations, Regional Office for Asia and the Pacific. ISBN 978-92-5-106488-7.
  36. ↑ Jharna Chakravorty, Sampat Ghosh, & Victor Benno Meyer-Rochow (2011). "Practices of entomophagy and entomotherapy by members of the Nyishi and Galo tribes, two ethnic groups of the state of Arunachal Pradesh (North-East India)". Journal of Ethnobiology and Ethnomedicine (BioMed Central) 7 (5). Retrieved April 30, 2011.
  37. ↑ Chen Shupei, Zhou Liang Yi. "荔枝椿象(Tessaratoma papillosa (Drury))" (in Chinese). Taiwan Agricultural Research Institute. Retrieved April 30, 2011.
  38. ↑ David A. Rider (October 5, 2009). "Plant Host Records: Tessaratomidae". Department of Entomology, North Dakota State University. Retrieved April 30, 2011.
  39. ↑ Subekti Rahayu, Koen Smets, Martin Lindgren & Fauzan Azhima (2005). "Hama repong damar dan pengendaliannya" (in Indonesian). World Agroforestry Centre - ICRAF, SEA Regional Office. Retrieved April 30, 2011.
  40. ↑ Bruce R. French. Insect Pests of Food Plants of Papua New Guinea: A compendium. Retrieved April 30, 2011.
  41. ↑ 41.0 41.1 L.H. Rolston, R.L. Aalbu, M.J. Murray, & D.A. Rider (1993). "A Catalog of the Tessaratomidae of the World". Papua New Guinea Journal of Agriculture, Forestry and Fisheries 36 (2): 36–108. Retrieved April 30, 2011.
  42. ↑ Robert G. Foottit, Peter Holdridge Adler (2009). Insect biodiversity: science and society. Cornell University Press. p. 247. ISBN 978-1-4051-5142-9.
  43. ↑ Philippe Magnien (October 31, 2009). "A catalog of Tessaratomidae". Heteroptera, un site pour les punaises. Retrieved April 30, 2011.
  44. ↑ David A. Rider (1998). "Nomenclatural Changes in the Pentatomoidea (Hemiptera-Heteroptera: Pentatomidae, Tessaratomidae). III. Generic Level Changes". Proc. Entomol. Soc. Wash. 100 (3): 504–510. Retrieved April 30, 2011.
  45. ↑ David A. Rider (September 18, 2009). "Genus Index - L". Department of Entomology, North Dakota State University. Retrieved April 30, 2011.
  46. ↑ T.D.A. Cockerell (1931). "Insects from the Miocene (Latah) of Washington". Annals of the Entomological Society of America (Entomological Society of America) 24 (2): 307–309. ISSN 1938-2901. Retrieved April 30, 2011.
  47. ↑ Dimitri Forero (March 13, 2009). "Pentatomoidea". Tree of Life web project. Retrieved April 28, 2011.

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