Heteropteran
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The Heteroptera are a group of about 40,000 species of insects in the order
Hemiptera Hemiptera (; ) is an order of insects, commonly called true bugs, comprising more than 80,000 species within groups such as the cicadas, aphids, planthoppers, leafhoppers, assassin bugs, bed bugs, and shield bugs. They range in size from ...
. They are sometimes called "true bugs", though that name more commonly refers to the Hemiptera as a whole. "Typical bugs" might be used as a more unequivocal alternative, since the heteropterans are most consistently and universally termed "bugs" among the Hemiptera. "Heteroptera" is Greek for "different wings": most species have
forewing Insect wings are adult outgrowths of the insect exoskeleton that enable insects to fly. They are found on the second and third thoracic segments (the mesothorax and metathorax), and the two pairs are often referred to as the forewings and hindwi ...
s with both membranous and hardened portions (called
hemelytra An elytron (; ; : elytra, ) is a modified, hardened forewing of beetles (Coleoptera), though a few of the true bugs (Hemiptera) such as the family Schizopteridae are extremely similar; in true bugs, the forewings are called hemelytra (sometimes ...
); members of the primitive sub-group
Enicocephalomorpha Enicocephaloidea is a sole superfamily within the infraorder Enicocephalomorpha of the hemipteran suborder Heteroptera ("true bugs"). The group was formerly thought to be related to the Reduviidae because of similarities in head structure but the ...
have completely membranous wings. The name "Heteroptera" is used in two very different ways in modern classifications. In Linnean nomenclature, it commonly appears as a
suborder Order () is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and recognized ...
within the
order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...
Hemiptera, where it can be
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
or
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
depending on its delimitation. In
phylogenetic nomenclature Phylogenetic nomenclature is a method of nomenclature for taxa in biology that uses phylogenetic definitions for taxon names as explained below. This contrasts with the traditional method, by which taxon names are defined by a '' type'', which c ...
, it is used as an unranked
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
within the
Prosorrhyncha The name Prosorrhyncha is a name (proposed by Sorensen et al. 1995) for a suborder of Hemiptera, comprising a grouping of the traditional taxon "Heteroptera" plus its sister taxon, the family Peloridiidae (often classified as a suborder itself). T ...
clade, which in turn is in the Hemiptera clade. This results from the realization that the
Coleorrhyncha Coleorrhyncha or Peloridiomorpha, also known as moss bugs or beetle bugs, are a suborder of Hemiptera and represent an ancient lineage of moss-feeding insects. They show some similarities to the Heteroptera but have been considered distinct. It h ...
are just "
living fossil A living fossil is a Deprecation, deprecated term for an extant taxon that phenotypically resembles related species known only from the fossil record. To be considered a living fossil, the fossil species must be old relative to the time of or ...
" relatives of the traditional Heteroptera, close enough to them to be united with that group. The infraorders
Leptopodomorpha Leptopodomorpha is an infraorder of insects in the order Hemiptera (true bugs). Leptopodomorpha is an infraorder of the order Heteroptera that contains more than 380 species. These small insects are also called shore bugs, or spiny shore bugs. As ...
,
Gerromorpha The Gerromorpha comprise an infraorder of insects in the "true bug" order (biology), order Hemiptera. These "typical" bugs (suborder Heteroptera) are commonly called semiaquatic bugs or shore-inhabiting bugs. The Ochteroidea (infraorder Nepomorph ...
, and
Nepomorpha Nepomorpha is an infraorder of insects in the "true bug" order (biology), order (Hemiptera). They belong to the "typical" bugs of the suborder Heteroptera. Due to their Aquatic insects, aquatic habits, these animals are known as true Heteroptera# ...
, comprise a significant component of the world's
aquatic Aquatic means relating to water; living in or near water or taking place in water; does not include groundwater, as "aquatic" implies an environment where plants and animals live. Aquatic(s) may also refer to: * Aquatic animal, either vertebrate ...
and semiaquatic insects. There are 23 families, 343 genera and 4,810 species group taxa within these three infraorders. Most of the remaining groups that are common and familiar are in the
Cimicomorpha The Cimicomorpha are an infraorder of insects in the order Hemiptera, the true bugs. The Rostrum (anatomy), rostrum and other Morphology (biology), morphology of some members apparently is adapted to feeding on animals as their prey or host (biol ...
and
Pentatomomorpha The Pentatomomorpha comprise an infraorder of insects in the true bug order Hemiptera. It unites such animals as the shield- or stink-bugs (Pentatomidae and alies), flat bugs (Aradidae), seed bugs (Lygaeidae and Rhyparochromidae), etc. They ar ...
.


Classification

The use of the name "Heteroptera" has had the rank of
order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...
, dating back to 1810 by
Pierre André Latreille Pierre André Latreille (; 29 November 1762 – 6 February 1833) was a French zoology, zoologist, specialising in arthropods. Having trained as a Roman Catholic priest before the French Revolution, Latreille was imprisoned, and only regained hi ...
. Only recently has it been relegated to a subsidiary rank within a larger definition of Hemiptera, so many reference works still include it as an order. Whether to continue treating it as a suborder is still a subject of some controversy, as is whether the name itself should ever be used, although three basic approaches ranging from abolishing it entirely to maintaining the
taxonomy image:Hierarchical clustering diagram.png, 280px, Generalized scheme of taxonomy Taxonomy is a practice and science concerned with classification or categorization. Typically, there are two parts to it: the development of an underlying scheme o ...
with a slight change in
systematics Systematics is the study of the diversification of living forms, both past and present, and the relationships among living things through time. Relationships are visualized as evolutionary trees (synonyms: phylogenetic trees, phylogenies). Phy ...
is proposed, two of which (but not the traditional one) agree with the
phylogeny A phylogenetic tree or phylogeny is a graphical representation which shows the evolutionary history between a set of species or Taxon, taxa during a specific time.Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, M ...
. The competing classifications call for a preference for two suborders versus one when the "living fossil" family
Peloridiidae The Peloridiidae or moss bugs are a family of true bugs, comprising eighteen genera and thirty-four species. They are small, ranging in length from 2 to 4 mm, rarely seen, peculiarly lumpy, flattened bugs found in Patagonia (Argentina and Chi ...
is taken into consideration: In one revised classification proposed in 1995, the name of the suborder is
Prosorrhyncha The name Prosorrhyncha is a name (proposed by Sorensen et al. 1995) for a suborder of Hemiptera, comprising a grouping of the traditional taxon "Heteroptera" plus its sister taxon, the family Peloridiidae (often classified as a suborder itself). T ...
, and Heteroptera is a rankless subgroup within it. The only difference between Heteroptera and Prosorrhyncha is that the latter includes the family
Peloridiidae The Peloridiidae or moss bugs are a family of true bugs, comprising eighteen genera and thirty-four species. They are small, ranging in length from 2 to 4 mm, rarely seen, peculiarly lumpy, flattened bugs found in Patagonia (Argentina and Chi ...
, which is a tiny relictual group that is in its own
monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unisp ...
superfamily and infraorder. In other words, the Heteroptera and Prosorrhyncha ''sensu'' Sorensen et al. are identical except that Prosorrhyncha contains one additional infraorder, called
Peloridiomorpha Coleorrhyncha or Peloridiomorpha, also known as moss bugs or beetle bugs, are a suborder of Hemiptera and represent an ancient lineage of moss-feeding insects. They show some similarities to the Heteroptera but have been considered distinct. It h ...
(comprising only 13 small genera). The ongoing conflict between traditional, Linnaean classifications and nontraditional classifications is exemplified by the problem inherent in continued usage of the name Heteroptera when it no longer can be matched to any standard Linnaean rank (as it falls below suborder but above infraorder). If this classification succeeds, then the "Heteroptera" grouping may be discarded, but in that case it is likely that no ranks will be used at all according to the standards of
phylogenetic nomenclature Phylogenetic nomenclature is a method of nomenclature for taxa in biology that uses phylogenetic definitions for taxon names as explained below. This contrasts with the traditional method, by which taxon names are defined by a '' type'', which c ...
. In the traditional classification, the Peloridiidae are retained as their own suborder, called
Coleorrhyncha Coleorrhyncha or Peloridiomorpha, also known as moss bugs or beetle bugs, are a suborder of Hemiptera and represent an ancient lineage of moss-feeding insects. They show some similarities to the Heteroptera but have been considered distinct. It h ...
; "Heteroptera" is treated the same. Functionally, the only difference between this classification and the preceding is that the former uses the name Prosorrhyncha to refer to a particular clade, while the traditional approach divides this into the
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
Heteroptera and the
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
Coleorrhyncha. Many believe it is preferable to use only one name because the characteristics of the two traditional suborders are too closely related to be treated as separate. Alternatively,Cassis, Gerasimos & Gross, Gordon (1995): Australian Biological Resources Study â€
Hemiptera: Heteroptera (Coleorrhyncha to Cimicomorpha). Gerrids, Reduviids, Water-striders
Version of June 30, 1995. Retrieved July 28, 2008.
the modified approach of placing Coleorrhyncha 'within' the Heteroptera can be used. Indeed, as that solution preserves the well-known Heteroptera at the taxonomic rank they traditionally hold while making them a good
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
group, it seems preferable to the paraphyletic "Heteroptera" used in older works. In that case, the "core" Heteroptera could be considered a
section Section, Sectioning, or Sectioned may refer to: Arts, entertainment and media * Section (music), a complete, but not independent, musical idea * Section (typography), a subdivision, especially of a chapter, in books and documents ** Section sig ...
– as yet unnamed, mainly because the Prosorrhyncha were proposed earlier – within the "expanded" Heteroptera, or the latter could simply be described as consisting of a basal "living fossil" lineage and a more
apomorph In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to ha ...
ic main radiation. Whether the name "Coleorrhyncha" is to be retained for the basal lineage or whether the more consistent "Peloridiomorpha" is used instead is a matter of taste, as described below. Separate from the question of the actual "closeness" of Heteroptera and Coleorrhyncha is the potential disruption to traditional construction of names; there seems to be reluctance among hemipterists to abandon the use of "Heteroptera". This can be seen by the name itself, as it is a violation of convention to use the ending "-ptera" for any rank above genus other than an order – though since it is a convention rather than a mandatory rule of Linnean nomenclature, taxonomists are technically free to violate it (which is why, for example, not all insect orders end in "-ptera", e.g.,
Odonata Odonata is an order of predatory flying insects that includes the dragonflies and damselflies (as well as the '' Epiophlebia'' damsel-dragonflies). The two major groups are distinguished with dragonflies (Anisoptera) usually being bulkier with ...
). However, in most cases when such conventions are violated, it does not create an internal conflict as in the present case (that is, the order Hemiptera has a suborder named Heteroptera, which is an internal conflict). At least some hemipterists argue that the name Heteroptera should be dropped entirely to eliminate this internal conflict, though the third possibility offers a workaround. In that case, to achieve full consistency of names "Coleorrhyncha" would probably be dropped in favor of "Peloridiomorpha".


References


Further reading

*


External links

*Heteropteran Systematics Lab at University of California, Riverside http://heteroptera.ucr.edu
Identification guide to UK Hemiptera
{{Authority control Hemiptera Insect suborders Extant Triassic first appearances