Tessaratomidae
Tessaratomidae | |
---|---|
Pycanum rubens, an adult from the Gunung Mulu National Park, Malaysia | |
Scientific classification | |
Domain: | Eukaryota |
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 | |
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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
All species are exclusively
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
The head of tessaratomids is generally small and triangular, with the
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
Some tessaratomids guard their eggs and nymphs from
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.[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]
Maternal care
In 1998, Gogala et al. described tessaratomines of the genus
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 onto 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.[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 cm (5.9 to 10.6 in).[21]
The chemicals produced by
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
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
Right: Parasitoid wasps of the genus Trissolcus emerging from the eggs of a pentatomid, Graphosoma italicum
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
Economic significance
As food
The edible stink bug
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
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,
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. |
Bronze orange bugs (Musgraveia sulciventris) are serious pests to
In
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
Tessaratomidae is classified under
They are mostly found in tropical Africa, Asia, and Oceania though a few species can be found in the
Listed below are the three subfamilies; their authors and type genera; the tribes, subtribes, and genera classified under them; and their
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[44]
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
- 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
- type genus: Tessaratoma Lepeletier & Serville, 1825
- Anacanthopus - Philippines
- Asiarcha - China, India, Indochina
- Aurungabada - India (Bombay)
- Candace - West Africa
- *Candace intermedia
- *Candace platygastra
- *Candace virescens
- 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 Tessaratomina 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
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]
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See also
References
- ISBN 978-0-306-44967-3.
- ^ Philippe Magnien (December 12, 2007). "Sepina aberrans (Schouteden, 1905)". Heteroptera, un site pour les punaises. Retrieved April 30, 2011.
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- ^ "Hou Zuki an" 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.
- ^ ISBN 978-1-77007-061-5.
- ^ ISBN 978-0-8014-2066-5.
- ^ a b c d 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)" (PDF). Taiwan Journal of Biodiversity (台灣生物多樣性研究) (in English and Chinese). 12 (4): 393–406. Retrieved April 30, 2011.
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- ^ . Retrieved May 1, 2011.
- ^ a b c d e f g h Philippe Magnien; Koen Smets; Dominique Pluot-Sigwalt & Jérôme Constant (2008). "A new species of Pygoplatys Dallas (Heteroptera, Tessaratomidae) from the Damar agroforests in Sumatra: description, immatures and biology" (PDF). Nouvelle Revue d'Entomologie. 24 (2): 99–112. Retrieved April 30, 2011.
- ^ a b Jonathan Figueroa Jiménez & Nohely Trabal. "Piezosternum subulatum (Thunberg 1783)" (PDF). Department of Biology, University of Puerto Rico at Mayagüez. Retrieved May 1, 2011.
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(help) - ^ ISBN 978-1-4020-6242-1.
- ^ a b c Tony, Sandy & Peter Chew (August 7, 2010). "Family Tessaratomidae - Large Stink Bugs". Brisbane Insects and Spiders. Retrieved May 1, 2011.
- ^ S. Tachikawa (1991). Studies on the Subsociality of Japanese Heteroptera(in Japanese). Tokyo Agricultural Press, Tokyo.
- ^ ISBN 978-0-521-58977-2.
- ISSN 1210-5759. Retrieved April 30, 2011.
- ^ S2CID 41951432. Retrieved April 27, 2011.
- ^ Kyle R. Conrad (2009). "Parental Care in Arthropods: Bracket Organization and Discussion of Behaviors" (PDF): 4. Retrieved April 30, 2011.
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(help) - ^ a b c d e Geoff B. Monteith (2011). "Maternal Care, Food Plants and Distribution of Australian Oncomerinae (Hemiptera: Heteroptera: Tessaratomidae)" (PDF). Australian Entomologist. 38 (1): 37–48. Archived from the original (PDF) on July 6, 2011. Retrieved April 30, 2011.
- ^ Ashraf M. El-Sayed (November 6, 2009). "Genus Compounds: Tessaratoma, Family: Tessaratomidae, Order: Heteroptera". The Pherobase. Retrieved May 1, 2011.
- ^ a b Jan Raška (2009). Function of metathoracic scent glands in terrestrial Heteroptera (PDF) (Bachelor thesis). Univerzita Karlova v Praze. Retrieved May 1, 2011.
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- ^ a b c David A. Rider (January 14, 2008). "Natural Enemies Index". Department of Entomology, North Dakota State University. Retrieved May 1, 2011.
- ISSN 1000-1034. Retrieved May 1, 2011.
- ^ "Parasitoids". Integrated Pest Management (IPM) Thailand. Retrieved May 5, 2011.
- ^ ISSN 0971-930X. Retrieved May 1, 2011.
- S2CID 84238759.
- ^ Cathy Dzerefos; David A. Rider (July 28, 2010). "Thongolifha (Encosternum delegorguei)". Department of Entomology, North Dakota State University. Retrieved April 30, 2011.
- ^ Cathy Dzerefos. "Traditional Cuisine With A Difference". Kruger National Park. Retrieved April 30, 2011.
- ISBN 978-0-7974-2897-3.
- ^ a b c Phillip Chidavaenzi (September 7, 2010). "Harurwa – more than food in Bikita". News Day. Retrieved April 30, 2011.
- ^ a b c Rob Toms & Mashudu Thagwana (2002). "Eat Your Bugs!". Science in Africa. Archived from the original on April 16, 2011. Retrieved April 30, 2011.
- ^ 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 (PDF) (Ph.D. thesis). University of Pretoria. Retrieved April 30, 2011.
- ISBN 978-92-5-106488-7.
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- ^ Chen Shupei; Zhou Liang Yi. "荔枝椿象(Tessaratoma papillosa (Drury))" (in Chinese). Taiwan Agricultural Research Institute. Retrieved April 30, 2011.
- ^ David A. Rider (October 5, 2009). "Plant Host Records: Tessaratomidae". Department of Entomology, North Dakota State University. Retrieved April 30, 2011.
- ^ 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.
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(help)CS1 maint: multiple names: authors list (link) - Bruce R. French. Insect Pests of Food Plants of Papua New Guinea: A compendium(PDF). Retrieved April 30, 2011.
- ^ a b L.H. Rolston; R.L. Aalbu; M.J. Murray & D.A. Rider (1993). "A Catalog of the Tessaratomidae of the World" (PDF). Papua New Guinea Journal of Agriculture, Forestry and Fisheries. 36 (2): 36–108. Retrieved April 30, 2011.
- ISBN 978-1-4051-5142-9.
- ^ Philippe Magnien (October 31, 2009). "A catalog of Tessaratomidae". Heteroptera, un site pour les punaises. Retrieved April 30, 2011.
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- ^ David A. Rider (1998). "Nomenclatural Changes in the Pentatomoidea (Hemiptera-Heteroptera: Pentatomidae, Tessaratomidae). III. Generic Level Changes" (PDF). Proc. Entomol. Soc. Wash. 100 (3): 504–510. Retrieved April 30, 2011.
- ^ David A. Rider (September 18, 2009). "Genus Index - L". Department of Entomology, North Dakota State University. Retrieved April 30, 2011.
- ISSN 1938-2901. Retrieved April 30, 2011.
- ^ Dimitri Forero (March 13, 2009). "Pentatomoidea". Tree of Life web project. Retrieved April 28, 2011.
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External links
- [1],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