Leaf Hoppers
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A leafhopper is the common name for any species from the family Cicadellidae. These minute insects, colloquially known as hoppers, are plant feeders that suck plant sap from grass, shrubs, or trees. Their hind legs are modified for jumping, and are covered with hairs that facilitate the spreading of a secretion over their bodies that acts as a water repellent and carrier of pheromones. They undergo a partial
metamorphosis Metamorphosis is a biological process by which an animal physically develops including birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation. Some inse ...
, and have various host associations, varying from very generalized to very specific. Some species have a cosmopolitan distribution, or occur throughout the temperate and tropical regions. Some are pests or vectors of plant viruses and phytoplasmas. The family is distributed all over the world, and constitutes the second-largest hemipteran family, with at least 20,000 described species. They belong to a lineage traditionally treated as infraorder Cicadomorpha in the
suborder Order ( la, ordo) 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 ...
Auchenorrhyncha, but as the latter taxon is probably not
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
, many modern authors prefer to abolish the Auchenorrhyncha and elevate the cicadomorphs to a suborder Clypeorrhyncha. Members of the tribe Proconiini of the
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end subfamily botanical names with "-oideae", and zoologi ...
Cicadellinae CicadellinaeLatreille (1825) is a leafhopper subfamily in the family Cicadellidae. Selected genera * '' Bothrogonia'' * ''Cicadella'' * ''Cofana'' * ''Graphocephala'' * ''Homalodisca'' * ''Zyzzogeton ''Zyzzogeton'' is a rare genus of leaf ...
are commonly known as sharpshooters.


Description and ecology

The Cicadellidae combine the following features: * The thickened part of the
antenna Antenna ( antennas or antennae) may refer to: Science and engineering * Antenna (radio), also known as an aerial, a transducer designed to transmit or receive electromagnetic (e.g., TV or radio) waves * Antennae Galaxies, the name of two collid ...
e is very short and ends with a bristle (arista). * Two ocelli (simple eyes) are present on the top or front of the head. * The tarsi are made of three segments. * The femora are at front with, at most, weak spines. * The hind tibiae have one or more distinct keels, with a row of movable spines on each, sometimes on enlarged bases. * The base of the middle legs is close together where they originate under the thorax. * The front wings not particularly thickened. An additional and unique character of leafhoppers is the production of
brochosome Brochosomes are intricately structured microscopic granules secreted by leafhoppers (the family Cicadellidae of the insect order Hemiptera) and typically found on their body surface and, more rarely, eggs. Brochosomes were first described in 1 ...
s, which are thought to protect the animals, and particularly their egg
clutch A clutch is a mechanical device that engages and disengages power transmission, especially from a drive shaft to a driven shaft. In the simplest application, clutches connect and disconnect two rotating shafts (drive shafts or line shafts). ...
es, from predation as well as pathogens. Like other
Exopterygota The Exopterygota (Ancient Greek ''ἔξω'' (éxō, “outside”) + ''πτερόν'' (pterón, “wing”) + New Latin ''-ota'' (“having”)), also known as Hemipterodea, are a superorder of insects of the subclass Pterygota in the infraclas ...
, the leafhoppers undergo direct development from nymph to adult without a pupal stage. While many leafhoppers are drab little insects as is typical for the Membracoidea, the adults and nymphs of some species are quite colorful. Some – in particular Stegelytrinae – have largely translucent wings and resemble flies at a casual glance. Leafhoppers have
piercing-sucking mouthpart Insects have mouthparts that may vary greatly across insect species, as they are adapted to particular modes of feeding. The earliest insects had chewing mouthparts. Most specialisation of mouthparts are for piercing and sucking, and this mode o ...
s, enabling them to feed on plant sap. A leafhoppers' diet commonly consists of sap from a wide and diverse range of plants, but some are more
host A host is a person responsible for guests at an event or for providing hospitality during it. Host may also refer to: Places * Host, Pennsylvania, a village in Berks County People *Jim Host (born 1937), American businessman * Michel Host ...
-specific. Leafhoppers mainly are
herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
s, but some are known to eat smaller insects, such as aphids, on occasion. A few species are known to be mud-puddling, but as it seems, females rarely engage in such behavior. Many species are also known to opportunistically pierce the human skin and draw blood but the function of such behaviour is unclear. Leafhoppers can transmit
plant pathogen Plant pathology (also phytopathology) is the scientific study of diseases in plants caused by pathogens (infectious organisms) and environmental conditions (physiological factors). Organisms that cause infectious disease include fungi, oomyc ...
s, such as viruses, phytoplasmas and bacteria. Cicadellidae species that are significant agricultural pests include the
beet leafhopper The beet leafhopper (''Circulifer tenellus''), also sometimes known as ''Neoaliturus tenellus'', is a species of leafhopper which belongs to the family Cicadellidae in the order Hemiptera. Morphology A lot of morphological diversity has be ...
(''Circulifer tenellus''), the maize leafhopper (''
Cicadulina mbila ''Cicadulina mbila'', the maize leafhopper, is a leafhopper species in the genus '' Cicadulina''. It is found in sub-Saharan Africa (Kenya, Tanzania and Uganda), the Middle East, West Asia and India. It is one of the vectors of the maize strea ...
''), potato leafhopper (''
Empoasca fabae ''Empoasca fabae'' belongs to family Cicadellidae and genus Empoasca within order Hemiptera.”Lamp, W., D., Miranda, L., Culler & Alexander, L. (2011) “Host Suitability and Gas Exchange Response of Grapevines to Potato Leafhopper (Hemiptera: ...
''), two-spotted leafhopper ('' Sophonia rufofascia''),
blue-green sharpshooter ''Graphocephala atropunctata'', commonly known as the blue-green sharpshooter, is a hemipteran bug endemism, endemic to California. It carries the phytopathogenic bacteria ''Xylella fastidiosa'' which infects the xylem of grape vines causing die- ...
('' Graphocephala atropunctata''),
glassy-winged sharpshooter The glassy-winged sharpshooter (''Homalodisca vitripennis'', formerly known as ''H. coagulata'') is a large leafhopper (family Cicadellidae), similar to other species of sharpshooter. Description These sharpshooters are about in length. Thei ...
(''Homalodisca vitripennis''), the
common brown leafhopper The common brown leafhopper, ''Orosius orientalis'' (Matsumura) ( Hemiptera: Cicadellidae) is one of the most common species of Australian leafhoppers with a very wide host range. It is an important vector of several viruses and phytoplasmas wo ...
(''Orosius orientalis''), rice green leafhoppers (''
Nephotettix ''Nephotettix'' is a genus of planthoppers in the subfamily Deltocephalinae and tribe Chiasmini. Species are mostly found in Asia, although two are from Africa. Known as 'green leafhoppers' of paddy/rice, ''Nephotettix cincticeps'' and ''N. vi ...
'' spp.), and the white apple leafhopper (''
Typhlocyba pomaria ''Typhlocyba'' is a genus of true bugs belonging to the family Cicadellidae. Synonym: ''Typhlocypha'' Scudder, 1881 Species: * ''Typhlocyba maderae'' * ''Typhlocyba quercus ''Typhlocyba'' is a genus of true bugs belonging to the family Cic ...
''). The
beet leafhopper The beet leafhopper (''Circulifer tenellus''), also sometimes known as ''Neoaliturus tenellus'', is a species of leafhopper which belongs to the family Cicadellidae in the order Hemiptera. Morphology A lot of morphological diversity has be ...
(''Circulifer tenellus'') can transmit the beet curly top virus to various members of the
nightshade family The Solanaceae , or nightshades, are a family of flowering plants that ranges from annual and perennial herbs to vines, lianas, epiphytes, shrubs, and trees, and includes a number of agricultural crops, medicinal plants, spices, weeds, and orna ...
, including tobacco, tomato, or eggplant, and is a serious vector of the disease in chili pepper in the Southwestern United States. In some cases, the plant pathogens distributed by leafhoppers are also pathogens of the insects themselves, and can replicate within the leafhoppers' salivary glands. Leafhoppers are also susceptible to various insect pathogens, including Dicistroviridae viruses, bacteria and fungi; numerous
parasitoid In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionarily stable str ...
s attack the eggs and the adults provide food for small
insectivore A robber fly eating a hoverfly An insectivore is a carnivorous animal or plant that eats insects. An alternative term is entomophage, which can also refer to the human practice of eating insects. The first vertebrate insectivores wer ...
s. Some species such as the Australian ''Kahaono montana'' even build silk nests under the leaves of trees they live in, to protect them from predators.


Systematics

In the now-obsolete classification that was used throughout much of the 20th century, the leafhoppers were part of the Homoptera, a
paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
assemblage uniting the basal lineages of
Hemiptera Hemiptera (; ) is an order (biology), order of insects, commonly called true bugs, comprising over 80,000 species within groups such as the cicadas, aphids, planthoppers, leafhoppers, Reduviidae, assassin bugs, Cimex, bed bugs, and shield bugs. ...
and ranked as
suborder Order ( la, ordo) 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 ...
. The splitting of the Homoptera is likely to be repeated for the Auchenorrhyncha for similar reasons, as the Auchenorrhyncha simply seem to group the moderately advanced Hemiptera, regardless of the fact the highly
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 have ...
ic Coleorrhyncha and
Heteroptera The Heteroptera are a group of about 40,000 species of insects in the order Hemiptera. 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 al ...
(typical bugs) evolved from auchenorrhynchans. Hence, a recent trend treats the most advanced hemipterans as three or four lineages, namely Archaeorrhyncha (Fulgoromorpha if included in Auchenorrhyncha), Coleorrhyncha and Heteroptera (sometimes united as Prosorrhyncha) and Clypeorrhyncha. Within the latter, the three traditional superfamiliesCercopoidea (froghoppers and spittlebugs), Cicadoidea (cicadas) and Membracoidea – appear to be
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
. The leafhoppers are the most
basal Basal or basilar is a term meaning ''base'', ''bottom'', or ''minimum''. Science * Basal (anatomy), an anatomical term of location for features associated with the base of an organism or structure * Basal (medicine), a minimal level that is nec ...
living lineage of Membracoidea, which otherwise include the families Aetalionidae (aetalionid treehoppers), Membracidae (typical treehoppers and thorn bugs),
Melizoderidae Melizoderidae is a family of tree-hoppers restricted to South America with only two genera, ''Melizoderes'' and ''Llanquihuea''. The nymphs of melizoderids have the tergum of the 9th segment concealing the anal opening from above. The frontoclype ...
, and
Myerslopiidae Myerslopiidae is a family of tree-hoppers which consist of about 20 species in three genera with a distribution limited to New Zealand and Chile. Adult hoppers have a strongly sclerotized body with elytra-like tegmina meeting along the median. Th ...
.


Subfamilies

The leafhoppers are divided into 25
subfamilies In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end subfamily botanical names with "-oideae", and zoologi ...
, which are listed here alphabetically, as too little is known about the family's internal phylogeny. *
Aphrodinae Aphrodinae is a subfamily of leafhoppers in the family Cicadellidae. There are about 7 genera and at least 20 described species in Aphrodinae. Genera These seven genera belong to the subfamily Aphrodinae: * ''Anoscopus'' Kirschbaum, 1858 * ''Ap ...
*
Bathysmatophorinae Bathysmatophorinae is a small Subfamily (biology), subfamily in the family Cicadellidae (leafhoppers). Description Bathysmatophorinae is a rare and primitive leafhopper subfamily, with a robust appearance and dull brown or grey colouration. They ...
*
Cicadellinae CicadellinaeLatreille (1825) is a leafhopper subfamily in the family Cicadellidae. Selected genera * '' Bothrogonia'' * ''Cicadella'' * ''Cofana'' * ''Graphocephala'' * ''Homalodisca'' * ''Zyzzogeton ''Zyzzogeton'' is a rare genus of leaf ...
*
Coelidiinae Coelidiinae is a subfamily of leafhoppers in the family Cicadellidae. There are at least 8 tribes, 108 genera, and over 900 species in Coelidiinae. Tribes The subfamily Coelidiinae is made up of these tribes. The number of genera and species ...
* Deltocephalinae *
Errhomeninae Errhomeninae is an insect subfamily of the family Cicadellidae, or the leafhoppers. It contains 3 tribes. Classification * Bathysmatophorini Anufriev, 1978 ** † ''Ambericarda'' Szwedo & Gębicki, 1998 ** ''Ankosus'' Oman & Musgrave, 1975 ** ...
*
Euacanthellinae Euacanthellinae is a subfamily of leafhoppers, native to Australia, New Caledonia and Madagascar, and adventive An introduced species, alien species, exotic species, adventive species, immigrant species, foreign species, non-indigenous speci ...
* Eurymelinae *
Evacanthinae Evacanthinae is a Subfamily (biology), subfamily in the family Cicadellidae (leafhoppers). Distribution Members of Evacanthinae are found worldwide and are on every continent except for Antarctica. Tribes and genera There are five Tribe (biolo ...
*
Hylicinae Hylicinae is a Subfamily (biology), subfamily in the family Cicadellidae (leafhoppers). Description Hylicine leafhoppers are moderately robust insects and are brownish to greyish in colouration. They feed on Dicotyledon trees and shrubs. Distri ...
*
Iassinae Iassinae is a subfamily of leafhoppers in the family Cicadellidae. Tribes and selected genera Batracomorphini Authority: Krishnankutty, Dietrich, Dai & Siddappaji, 2016 * '' Batracomorphus'' Lewis, 1834 Bythoniini Authority: Linnavuori, 19 ...
* Jascopinae *
Ledrinae Ledrinae is a relatively small subfamily within the very large and diverse leafhopper family Cicadellidae. Originally placed in its own family, the "Ledridae", it is based on the type genus ''Ledra''. Description The Ledrinae are mostly g ...
*
Megophthalminae Megophthalminae is a subfamily of leafhopper A leafhopper is the common name for any species from the family Cicadellidae. These minute insects, colloquially known as hoppers, are plant feeders that suck plant sap from grass, shrubs, or trees. ...
*
Mileewinae Mileewinae is a small Subfamily (biology), subfamily in the family Cicadellidae (leafhoppers). It is closely related to Typhlocybinae and contains species that were previously part of Cicadellinae. Description Mileewinae leafhoppers are small, s ...
* Nastlopiinae *
Neobalinae Neobalinae is a small Subfamily (biology), subfamily in the family Cicadellidae (leafhoppers). Description Neobaline leafhoppers are often darkly coloured with fluorescent accents. Distribution Members of the subfamily are found feeding on Dico ...
*
Neocoelidiinae Neocoelidiinae is a small Subfamily (biology), subfamily in the family Cicadellidae (leafhoppers). It was originally included within the subfamily Coelidiinae. Description Neocoelidiinae are easily recognisable by their very long antennae and di ...
* Nioniinae * Phereurhininae * Portaninae * Signoretiinae * Tartessinae *
Typhlocybinae Typhlocybinae is a subfamily of insects in the leafhopper family, Cicadellidae. This is currently the second largest leafhopper subfamily based on the number of described species, but researchers believe there are so many taxa yet undescribed th ...
*
Ulopinae Ulopinae is a subfamily of leafhoppers. Tribes * Cephalelini * Coloborrhinini * Mesargini * Monteithiini * Ulopini Description Leafhoppers from this group have hardened, elytra An elytron (; ; , ) is a modified, hardened forewing of beet ...
* Further information: ''
Utecha trivia ''Utecha trivia'' is a European species of leafhopper. Description The species are in length. The males have a distinctive mark and a light yellowish colour base with black streaks starting from clavus (insect wing), clavus to the forewing. Th ...
''


See also

*
Planthopper A planthopper is any insect in the infraorder Fulgoromorpha, in the suborder Auchenorrhyncha, a group exceeding 12,500 described species worldwide. The name comes from their remarkable resemblance to leaves and other plants of their environment ...
* Treehopper *
List of animals that produce silk Silk is produced by a variety of animals, for different purposes, with various types being produced. Insects *Silkworms produce silk when undergoing larval to adult metamorphosis. This includes not only the domesticated ''Bombyx mori'', but a wide ...


References


Further reading

*Carver, M, FG. Gross, and TE. Woodward. 1991. Hemiptera (bugs, leafhoppers, cicadas, aphids, scale insects, etc.) In: ''The Insects of Australia – a Textbook for Students and Research Workers Volume 1.'' Melbourne University Press, Melbourne, Australia".


External links


Red-banded leafhopper ''Graphocephala coccinea'' – diagnostic photographs and information

Sharpshooter Leafhoppers of the World (Hemiptera: Cicadellinae) – Online Database with color photos of nearly all described species.


*



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