Scutelleridae

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Jewel bugs
Metallic shield bug444.jpg
Scutiphora pedicellata , a jewel bug from Australia
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hemiptera
Suborder: Heteroptera
Infraorder: Pentatomomorpha
Superfamily: Pentatomoidea
Family: Scutelleridae
Leach, 1815
Subfamilies

Scutelleridae is a family of true bugs. They are commonly known as jewel bugs or metallic shield bugs due to their often brilliant coloration. With the name based on the Asian genus Scutellera , they are also known as shield-backed bugs due to the enlargement of the thoracic scutellum into a continuous shield over the abdomen and wings. [1] This latter characteristic distinguishes them from most other families within Heteroptera, and may lead to misidentification as a beetle rather than a bug. These insects feed on plant juices from a variety of different species, including some commercial crops. Closely related to stink bugs, they may also produce an offensive odour when disturbed. There are around 450 species worldwide. [2]

Contents

Description

Jewel bugs are small to medium-sized oval-shaped bugs with a body length averaging at 5 to 20 mm (0.20 to 0.79 in). [3] They can easily be distinguished from stink bugs (Pentatomidae) because the shield-like enlarged last section of their thorax (known as the scutellum, Latin for "little shield") completely covers the abdomen and the wings. [4]

Despite their resemblance to beetles, jewel bugs are hemipterans or true bugs. [5] The scutellum is an extension of the thorax, unlike the elytra of beetles which are hardened forewings. As such, jewel bugs have four membranous wings underneath the scutellum in contrast to two in beetles. [6] The scutellum in jewel bugs also does not have a division in the middle and thus does not 'split open' when they take flight like in beetles. [7]

The heads of jewel bugs are triangular and the antennae have three to five segments. [8] Like all heteropterans, jewel bugs are characterized by a segmented beak-like mouthpart (known as the rostrum). [2] During feeding, jewel bugs inject proteolytic enzymes in their saliva into plants, digesting plant matter into a liquid form which they then suck up. [9] The tarsus has three segments (tarsomeres). [6]

Colours

Though some species are quite drab, [10] the most conspicuous jewel bugs are often brilliantly colored, exhibiting a wide range of iridescent metallic hues that change with the view angle. [6] The colors are the result of a combination of factors. Some species like Chrysocoris stockerus and Scutellera nobilis display colors from multiple thin layers of pigmented chitin. The colors often change or become duller when the specimens are dried, due to the topmost chitinous layer becoming opaque and obscuring the colors of the bottom layer. The colors can be restored by moistening the surfaces with water. [11]

Iridescence (or goniochromism) in jewel bugs like Poecilocoris lewisi are the result of structural coloration. Instead of pigments, the colors are caused by the interference, diffraction, or scattering of light by numerous tiny structures.[ citation needed ]

In Poecilocoris lewisi, multiple tiny conical protuberances around 900 nm in height and averaging at a diameter of 360 nm are scattered on the epicuticle. These structures affect light passing through them, producing their oily-looking blue sheen (known as the Tyndall effect or Mie scattering). [11]

In other species like the African shield bug ( Calidea panaethiopica ), the dorsal cuticle is dotted with tiny regularly spaced hemispherical cavities. The depressions act like Bragg mirrors. When light hits the pitted surface, it gives off multiple reflections resulting in the distinctive two tone yellow-blue iridescence. [12]

The colors and patterns on jewel bugs can vary significantly between instars and even within adults of a species. [13]

Jewel bugs are also known to mimic the colors, patterns, and shape of other organisms for defensive purposes. An example is the yellow-spotted black Steganocerus multipunctatus which exhibits Müllerian mimicry with the tortoise beetle Chiridopsis suffriani . [14]

Ecology and life cycle

All jewel bugs feed on plants (phytophagous). The eggs are laid in compact clusters. [15] They may be round or barrel-shaped with a lid or a cap at the top (known as the operculum). They also contain a ring of small protuberances near the cap called micropylar processes. They permit the passage of sperm into the egg for fertilization and enable gaseous exchange from within the egg and the outside world for the embryos. The eggs are white or cream colored when freshly laid but can change color as the embryo matures. [16] When hatching, the prolarva (the advanced embryo) exit the egg by opening the lid through peristaltic movements and with the help of a T-shaped internal structure in the egg (known as the egg burster). [16] [17]

Like all hemipterans, jewel bugs undergo incomplete metamorphosis (hemimetaboly) and do not possess larval and pupal stages. Instead the adults develop from several stages (instars) of nymphs (usually five) through successive moltings (ecdysis). Nymphs resemble the adults except for size and the absence of wings. They can be of different coloration or patterns from adults. [15]

Some species are known to exhibit parental care of eggs and nymphs. Notable examples of which are Cantao parentum , [18] Pachycoris klugii , [19] Pachycoris stalii , [20] Pachycoris torridus , and Tectocoris diophthalmus . [13] [21]

Reproduction

Chemical secretions from dorsal abdominal or sternal exocrine glands are used to attract mates by certain species of jewel bugs. [22] In certain genera (like Tectocoris , Psacasta , Odontoscelis , and Irochrotus ), males possess special unicellular glands in the abdomen known as the androconia (singular: androconium). They release sex pheromones when ruptured. [6]

Females possess a spermatheca, an ectodermal gland which opens into the oviduct. These serve as storage for sperm deposited by males. It contains glands which can nourish the spermatozoa until they can be released to fertilize eggs. [17]

Male jewel bugs of the genus Hotea possess an unusually large, spiky, and heavily sclerotized genitalia. They are used in a mating practice known as traumatic insemination, a result of evolutionary sexual conflict. Male Hotea bugs tear through the female reproductive ducts to deposit sperm, inflicting substantial damage to the female in the process. [23]

Defenses

Like stink bugs, a vast majority of jewel bugs, both adults and nymphs, are also capable of releasing pungent defensive chemicals from glands located on the sides of the thorax. [2] [7] Typical compounds exuded by jewel bugs include alcohols, aldehydes, and esters. [24]

Nymphs and adults often exhibit clustering behavior, being found in large numbers close to each other. This behavior is thought to have an evolutionary advantage. The more individuals present in an area, the stronger the odor of the chemicals released when the bugs are threatened. [18] If this fails, stink bugs will react to threat by flying away or dropping to the ground. [2]

Classification and evolution

Scutellerids were first described by the English zoologist William Elford Leach in 1815. [25] [26] It belongs to the order Hemiptera (true bugs), under the suborder Heteroptera and infraorder Pentatomomorpha. They are classified under the superfamily Pentatomoidea. They were formerly classified as a subfamily of Pentatomidae by George Willis Kirkaldy in 1909. [27] The earliest attempt to restore them to family status was in 1917 by Edward Payson Van Duzee. [28] Most authorities today regard it as a valid family group. [27] In phylogenetic studies in 2008 by Grazia et al., Scutelleridae was shown to be consistently monophyletic, basal to Acanthosomatidae, and distal to Plataspididae and Parastrachiidae. [28] [29] Below is the morphological unweighted tree of the superfamily Pentatomoidea after Grazia et al. (2008). [29]

      ←      
             

Urostylididae

             
             

Saileriolidae

             
             

Acanthosomatidae

             
             
             

Tessaratomidae

             

Dinidoridae

   Cydnidae   sensu lato   
             

Cydnidae

             

Thaumastellidae

             

Parastrachiinae

             
             

Thyreocoridae

             

Lestoniidae

             

Phloeidae

             
             

Scutelleridae

             

Plataspididae

             

Pentatomidae

             

Canopidae

             

Megarididae

Subfamilies and genera

The family is composed of about 81 genera and around 450 species worldwide. While the tribal and subfamilial classifications remain unclear, [30] they are divided into eight subfamilies sensu lato : Elvisurinae, Eurygastrinae, Hoteinae (sometimes classified under Pachycorinae), Odontoscelinae, Odontotarsinae, Pachycorinae, Scutellerinae, and Tectocorinae. [31] [32]

Elvisurinae

Auth.: Stål, 1872. Widespread distribution, with the type genus Elvisura found in southern Africa; only one species of Elvisurinae, Solenosthedium bilunatum , is found in Europe. [33]

  1. Austrotichus Gross, 1975 - monotypic A. rugosusGross, 1975
  2. Coleotichus White, 1839
  3. Elvisura Spinola, 1837
  4. Solenosthedium Spinola, 1837
  5. Solenotichus Martin, 1897

Eurygastrinae

Auth.: Amyot & Audinet-Serville, 1843

Tribe Eurygastrini Amyot & Audinet-Serville, 1843
  1. Eurygaster Laporte, 1833
  2. Polyphyma Jakovlev, 1877
Tribe Psacastini Mulsant & Rey, 1865
  1. Ceratocranum Reuter, 1890
  2. Periphima Jakovlev, 1889
  3. Periphymopsis Schouteden, 1904
  4. Promecocoris Puton, 1886
  5. Psacasta Germar, 1839
  6. Xerobia Stål, 1873

Hoteinae

Auth.: Carapezza, 2008

  1. Deroplax Mayr, 1864
  2. Ellipsocoris Mayr, 1864
  3. Hotea Amyot & Serville, 1843

Odontoscelinae

Auth.: Amyot & Audinet-Serville, 1843; mostly European

  1. Holonotellus Horváth, 1895
  2. Irochrotus Amyot & Serville, 1843
  3. Odontoscelis Laporte, 1833

Odontotarsinae

Auth.: Mulsant & Rey, 1865

Tribe Odontotarsini Mulsant & Rey, 1865
  1. Ahmadocoris Carapezza, 2009
  2. Alphocoris Germar, 1839
  3. Melanodema Jakovlev, 1880
  4. Odontotarsiellus Hoberlandt, 1955
  5. Odontotarsus Laporte, 1833
  6. Urothyreus Horváth, 1921
Tribe Phimoderini Fuente, 1974
  1. Euptychodera Bergroth, 1908
  2. Fokkeria Schouteden, 1904
  3. Morbora Distant, 1899
  4. Phimodera Germar, 1839
  5. Vanduzeeina Schouteden, 1904

Pachycorinae

Auth.: Amyot & Audinet-Serville, 1843

  1. Acantholomidea Sailer, 1945
  2. Agonosoma Laporte, 1833
  3. Ascanius (bug) Stål, 1868
  4. Brailovskylus Eger, 2017
  5. Camirus Stål, 1862
  6. Chelycoris Bergroth, 1891
  7. Chelyschema Bergroth, 1891
  8. Coptochilus Amyot & Serville, 1843
  9. Crathis (bug) Stål, 1861
  10. Diolcus Mayr, 1864
  11. Dystus Stål, 1862
  12. Ephynes Stål, 1868
  13. Galeacius Distant, 1889
  14. Homaemus Dallas, 1851
  15. Lobothyreus Mayr, 1864
  16. Misippus Stål, 1868
  17. Nesogenes Horváth, 1921
  18. Orsilochides Kirkaldy, 1909
  19. Pachycoris Burmeister, 1835
  20. Polytes Stål, 1868
  21. Sphyrocoris Mayr, 1864
  22. Stethaulax Bergroth, 1891
  23. Symphylus Dallas, 1851
  24. Testrina Walker, 1867
  25. Tetyra Fabricius, 1803
  26. Tiridates (bug) Stål, 1868

Scutellerinae

Auth.: Leach, 1815. Selected genera:

Tectocorinae McDonald & Cassis, 1984 (monotypic)

Economic significance

Eurygaster-cf-testudinaria-13-VII-2007-478.jpg
Odontotarsus grammicus.jpg
Cotton Harlequin Bugs.jpg
Left: Eurygaster testudinaria , a sunn pest from Germany. Center: Odontotarsus grammicus , another sunn pest from Spain. Right: An adult and nymph cotton harlequin bug ( Tectocoris diophthalmus ).

Though most jewel bugs do little harm to crop plants, [34] a few members of Scutelleridae are considered major agricultural pests. Together with some species of stink bugs, they are collectively known as sunn pests (also spelled as senn, soun, or shüne pests) or wheat bugs. [30] The most economically important species of which are members of the genus Eurygaster .

Eurygaster integriceps , in particular, is a very destructive pest of cereal crops in North Africa, the Balkans, and western and central Asia. Other scutellerids known under the name 'sunn pest' include members of the genus Odontotarsus , among others. [35] Methods of control for sunn pests have included biological pest control, using wasps of the family Scelionidae from the genera Trissolcus and Ooencyrtus .[ citation needed ]

The cotton harlequin bug ( Tectocoris diophthalmus ) is also an important pest of cotton crops and Hibiscus . [34] [36]

Conservation

Biological methods of pest control have sometimes backfired. A parasitoid fly which preys on hemipterans, Trichopoda pennipes was introduced to Hawaii to control the invasive species Nezara viridula , the southern green stink bug. The fly now threatens native species of bugs in Hawaii as well, particularly the Koa bug (Coleotichus blackburniae, a jewel bug species notable for not possessing stink glands) which has now become rare. [37]

See also

Related Research Articles

<span class="mw-page-title-main">Pentatomoidea</span> Superfamily of true bugs

The Pentatomoidea are a superfamily of insects in the suborder Heteroptera of the order Hemiptera. As hemipterans, they possess a common arrangement of sucking mouthparts. The roughly 7000 species under Pentatomoidea are divided into 21 families. Among these are the stink bugs and shield bugs, jewel bugs, giant shield bugs, and burrower bugs.

<span class="mw-page-title-main">Pentatomidae</span> Family of insects

Pentatomidae is a family of insects belonging to the order Hemiptera, generally called shield bugs or stink bugs. Pentatomidae is the largest family in the superfamily Pentatomoidea, and contains around 900 genera and over 4700 species. As hemipterans, the pentatomids have piercing sucking mouthparts, and most are phytophagous, including several species which are severe pests on agricultural crops. However, some species, particularly in the subfamily Asopinae, are predatory and may be considered beneficial.

<span class="mw-page-title-main">Pyrrhocoridae</span> Family of true bugs

Pyrrhocoridae is a family of insects with more than 300 species world-wide. Many are red coloured and are known as red bugs and some species are called cotton stainers because their feeding activities leave an indelible yellow-brownish stain on cotton crops. A common species in parts of Europe is the firebug, and its genus name Pyrrhocoris and the family name are derived from the Greek roots for fire "pyrrho-" and bug "coris". Members of this family are often confused with, but can be quickly separated from, Lygaeidae by the lack of ocelli on the top of the head.

<span class="mw-page-title-main">Coreidae</span> Family of insects

Coreidae is a large family of predominantly sap-sucking insects in the Hemipteran suborder Heteroptera. The name "Coreidae" derives from the genus Coreus, which derives from the Ancient Greek κόρις (kóris) meaning bedbug.

<span class="mw-page-title-main">Pentatominae</span> Subfamily of true bugs

Pentatominae is a subfamily of Pentatomidae, a family of shield bugs. This subfamily is the largest one within the Pentatomidae, having 4937 species classified in 938 genera. Species in this subfamily are phytophages and several of them are considered agricultural pests. Some invasive pentatomines such as Halyomorpha halys and Bagrada hilaris have been considered household pests. Higher systematics of the group have been revised by Rider et al.

<span class="mw-page-title-main">Alydidae</span> Family of true bugs

Alydidae, commonly known as broad-headed bugs, is a family of true bugs very similar to the closely related Coreidae. There are at least 60 genera and 300 species altogether. Distributed in the temperate and warmer regions of the Earth, most are tropical and subtropical animals; for example Europe has a mere 10 species, and only 2 of these occur outside the Mediterranean region.

<span class="mw-page-title-main">Harpactorinae</span> Subfamily of true bugs

The Harpactorinae are a large subfamily of the Reduviidae. About 300 genera and 2,000 species worldwide have been described. Some of the species of the genera Zelus, Pselliopus, Sinea, and Apiomerus are of interest as biological pest control agents.

<i>Calliphara nobilis</i> Species of jewel bug

Calliphara nobilis is a species of jewel bug found in Asia. Like all species of jewel bugs, it is phytophagous, feeding on the leaves, fruit and seeds of its host plants. This insect is notable for its multiple defense mechanisms: it is highly mobile and swarms disperse with a loud buzz when disturbed; it is aposematically colored, which serves as a warning to any would-be predators that it is unpalatable; and it possesses a robust chemical defense mechanism: it can secrete an irritating and toxic fluid from a pair of metathoracic scent glands when threatened.

<span class="mw-page-title-main">Scutellerinae</span> Subfamily of true bugs

Scutellerinae is a subfamily of shield-backed bugs erected by Leach in 1815.

<i>Cantao</i> (bug) Genus of true bugs

Cantao is a genus of true bugs in family Scutelleridae and tribe Scutellerini.

<span class="mw-page-title-main">Tessaratomidae</span> Family of true bugs

Tessaratomidae is a family of true bugs. It contains about 240 species of large bugs divided into 3 subfamilies and 56 genera.

<span class="mw-page-title-main">Thyreocoridae</span> Family of true bugs

The Thyreocoridae are a family of shield bugs, known by common names that include negro bugs or ebony bugs. Historically, a few authors have called this family "Corimelaenidae" (e.g.), but the name Thyreocoridae, published in 1843, has nomenclatural priority over Corimelaenidae, published in 1872. Other classifications have placed them as a subfamily within the broad family Cydnidae.

<span class="mw-page-title-main">Dinidoridae</span> Family of true bugs

Dinidoridae is a small family of hemipteran "true bugs" comprising about sixteen genera and a hundred species the Hemiptera suborder Heteroptera. As a group the family does not have any common name. Until the late 19th century they were generally regarded as a subfamily of Pentatomidae.

<i>Sphaerocoris annulus</i> Species of true bug

Sphaerocoris annulus, common names Picasso bug or Zulu hud bug, is a species of shield-backed bugs of the family Scutelleridae.

<span class="mw-page-title-main">Pentatomini</span> Tribe of true bugs

Pentatomini is a tribe of shield bugs in the subfamily of Pentatominae.

<span class="mw-page-title-main">Cydninae</span> Subfamily of true bugs

Cydninae is a subfamily of burrowing bugs in the family Cydnidae. There are about 11 genera and at least 40 described species in Cydninae.

<span class="mw-page-title-main">Carpocorini</span> Tribe of true bugs

Carpocorini is a tribe of stink bugs in the family Pentatomidae. There are more than 100 genera in Carpocorini.

<span class="mw-page-title-main">Phyllocephalinae</span> Subfamily of true bugs

The Phyllocephalinae are a subfamily of shield bugs erected by Amyot and Serville in 1843.

<span class="mw-page-title-main">Discocephalinae</span> Subfamily of true bugs

The Discocephalinae are a subfamily of shield bugs, erected by Fieber in 1860, and found mostly in the Americas.

References

  1. ↑ "Scutelleridae - Jewel Bugs". Commonwealth Scientific and Industrial Research Organisation (CSIRO) Entomology. Retrieved April 27, 2011.
  2. 1 2 3 4 John L. Capinera (2008). Encyclopedia of entomology. Springer. pp. 608–609. ISBN   978-1-4020-6242-1.
  3. ↑ Robert G. Foottit & Peter Holdridge Adler (2009). Insect biodiversity: science and society. John Wiley and Sons. p. 247. ISBN   978-1-4051-5142-9.
  4. ↑ Mike Picker; Charles Griffiths & Alan Weaving (2004). Field guide to insects of South Africa. Field Guide Series. Struik. p. 132. ISBN   978-1-77007-061-5.[ permanent dead link ]
  5. ↑ "Scutelleridae". National Center for Biotechnology Information (NCBI).
  6. 1 2 3 4 Randall T. Schuh & James Alexander Slater (1995). True bugs of the world (Hemiptera:Heteroptera): classification and natural history. Cornell University Press. ISBN   978-0-8014-2066-5.
  7. 1 2 "Family Scutelleridae - Shield-backed Bugs". BugGuide. Retrieved April 27, 2011.
  8. ↑ "Jewel bugs, Shield Backed Bugs -Family Scutelleridae". Brisbane Insects and Spiders Home Page. Retrieved April 27, 2011.
  9. ↑ E. Grünbacher & B. Kromp (2008). "Investigations on the occurrence of wheat bugs (Scutelleridae, Pentatomidae; Heteroptera) in organic farming of Eastern Austria" (PDF). 1st Scientific Conference within the framework of the 8th European Summer Academy on Organic Farming. Retrieved April 27, 2011.{{cite journal}}: Cite journal requires |journal= (help)
  10. ↑ Scott Fabricant, Mariella Herberstein & Darrell Kemp. "Mechanisms and Functions of Colour in the Hibiscus Harlequin Bug" (PDF). The Australasian Society for the Study of Animal Behaviour. Retrieved April 28, 2011.{{cite journal}}: Cite journal requires |journal= (help)
  11. 1 2 Shūichi Kinoshita (2008). Structural colors in the realm of nature. World Scientific. p. 155. ISBN   978-981-270-783-3.
  12. ↑ Jean-Pol Vigneron, Moussa Ouedraogo, Jean-François Colomer, and Marie Rassart (200). "Spectral sideband produced by a hemispherical concave multilayer on the African shield-bug Calidea panaethiopica (Scutelleridae)". Physical Review E. 79 (2). American Physical Society: 021907. doi:10.1103/PhysRevE.79.021907. PMID   19391778.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  13. 1 2 Luis Cervantes Peredo (2002). "Description, Biology, and Maternal Care of Pachycoris klugii (Heteroptera: Scutelleridae)" (PDF). Florida Entomologist. 85 (3). Instituto de Ecologia, Florida Center for Library Automation: 464–473. doi: 10.1653/0015-4040(2002)085[0464:dbamco]2.0.co;2 . Retrieved April 27, 2011.
  14. ↑ Hugh D.C. Heron (2009). "Polymorphism in four tortoise beetles from Queensburgh, South Africa (Chrysomelidae: Cassidinae)" (PDF). Genus. 22 (1). University of Wrocław: 133–149. Retrieved April 28, 2011.
  15. 1 2 Maurice Burton & Robert Burton (1970). The international wildlife encyclopedia. Vol. 1. Marshall Cavendish. pp. 2344–2345. ISBN   978-0-7614-7266-7.
  16. 1 2 Selami Candan, Zekiye Suludere & Dilek Durak (2005). "Ultrastructure of the Eggs Chorion of Ceraleptus obtusus (Brulle, 1839)(Heteroptera: Coreidae)". Ohio Journal of Science. 105 (5). The Knowledge Bank, Ohio State University: 138–141. hdl:1811/31918.
  17. 1 2 Selami Candan, Zekiye Suludere & Mahmut Erbey (2008). "Morphology of eggs and spermatheca of Odontotarsus purpureolineatus (Heteroptera, Scutelleridae)". Biologia, Bratislava. 62 (6). Springer: 763–769. doi: 10.2478/s11756-007-0137-x . ISSN   0006-3088.
  18. 1 2 "Scutelleridae: Shield-backed Bugs". Save Our Waterways Now . Retrieved April 28, 2011.
  19. ↑ James T. Costa (2006). The other insect societies. Harvard University Press. p. 281. ISBN   978-0-674-02163-1.
  20. ↑ Livy Williams III, Maria C Coscarón, Pablo M Dellapé, and Timberley M Roane (2005). "The shield-backed bug, Pachycoris stallii: Description of immature stages, effect of maternal care on nymphs, and notes on life history". J Insect Sci. 5 (29). University of Wisconsin Libraries: 29. doi:10.1093/jis/5.1.29. PMC   1615236 . PMID   17119611.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  21. ↑ David A. Rider. "Pentatomoid Species Known to Exhibit Parental Care". Department of Entomology, North Dakota State University. Retrieved April 27, 2011.
  22. ↑ Ring T. Cardé & William J. Bell (1995). Chemical ecology of insects 2. Springer. p. 336. ISBN   978-0-412-03961-4.
  23. ↑ Göran Arnqvist & Locke Rowe (2005). Sexual conflict. Princeton University Press. p. 129. ISBN   978-0-691-12218-2.
  24. ↑ Stefan Schulz (2005). The chemistry of pheromones and other semiochemicals. Vol. 2. Springer. p. 37. ISBN   978-3-540-21308-6.
  25. ↑ "Scutelleridae". Integrated Taxonomic Information System.
  26. ↑ "Scutelleridae Leach, 1815". The Encyclopedia of Life .
  27. 1 2 G. Cassis, Australia. Bureau of Flora and Fauna, & Gordon F. Gross (2002). Zoological catalogue of Australia: Hemiptera: Heteroptera (Pentatomomorpha). CSIRO Publishing. p. 353. ISBN   978-0-643-06875-9.{{cite book}}: CS1 maint: multiple names: authors list (link)
  28. 1 2 Jocelia Grazia; Randall T. Schuhb & Ward C. Wheeler (2008). "Phylogenetic relationships of family groups in Pentatomoidea based on morphology and DNA sequences (Insecta: Heteroptera)" (PDF). Cladistics. 24 (6). Wiley-Blackwell: 932–976. doi:10.1111/j.1096-0031.2008.00224.x. PMID   34892882. S2CID   41951432 . Retrieved April 27, 2011.
  29. 1 2 Dimitri Forero (March 13, 2009). "Pentatomoidea". Tree of Life Web Project . Retrieved April 28, 2011.
  30. 1 2 Carl W . Schaefer; John D . Lattin & M . Javahery (2001). "Shield Bugs (Scutelleridae)". In Carl W . Schaefer & Antonio Ricardo Panizzi (eds.). Heteroptera of Economic Importance. CRC Press. pp. 475–503. doi:10.1201/9781420041859.ch14 (inactive 2024-11-12). ISBN   978-1-4200-4185-9.{{cite book}}: CS1 maint: DOI inactive as of November 2024 (link)
  31. ↑ Biolib: family of shield-backed bugs, Scutelleridae Leach, 1815 (retrieved 10 February 2024)
  32. ↑ David A. Rider. "Classification". Department of Entomology, North Dakota State University. Retrieved April 27, 2011.
  33. ↑ Škorput, Jadranka; Martinović, Matea; van der Heyden, Torsten; Skejo, Josip (March 2020). "Solenosthedium bilunatum (Heteroptera: Scutelleridae) at the Adriatic Coast of Croatia". ResearchGate. Entomologie heute 31 (2019): 25-29. Retrieved 4 February 2024.
  34. 1 2 Murray Fletcher (2007). "Plant bugs" (PDF). Primefact. 508. NSW Department of Primary Industries: 1–4. Retrieved April 27, 2011.
  35. ↑ Randall T. Schuh & James Alexander Slater (1995). True bugs of the world (Hemiptera:Heteroptera): classification and natural history. Cornell University Press. p. 36. ISBN   978-0-8014-2066-5.
  36. ↑ Peter T. Bailey (2007). Pests of field crops and pastures: identification and control. CSIRO Publishing. pp. 97–98. ISBN   978-0-643-06758-5.
  37. ↑ Roy Van Driesche, Mark Hoddle & Ted Center (2008). Control of pests and weeds by natural enemies: an introduction to biological control. John Wiley and Sons. ISBN   978-1-4051-4571-8.
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