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Treehoppers (more precisely typical treehoppers to distinguish them from the
Aetalionidae Aetalionidae are a family of treehoppers in the superfamily Membracoidea The superfamily Membracoidea of sap-sucking true-bugs includes two of the largest families within what used to be called the "Homoptera": the leafhoppers ( Cicadellidae) ...
) and thorn bugs are members of the
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
Membracidae, a group of
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s related to the
cicada The cicadas () are a superfamily, the Cicadoidea, of insects in the order Hemiptera (true bugs). They are in the suborder Auchenorrhyncha, along with smaller jumping bugs such as leafhoppers and froghoppers. The superfamily is divided into two ...
s and the
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. Their hind legs are modified for jumping, and a ...
s. About 3,200
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
of treehoppers in over 400
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
are known.Treehoppers.
Dr. Metcalf. NCSU Libraries. North Carolina State University.
They are found on all continents except Antarctica; only five species are known from
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
. Individual treehoppers usually live for only a few months.


Morphology

Treehoppers, due to their unusual appearance, have long interested naturalists. They are best known for their enlarged and ornate
pronotum The prothorax is the foremost of the three segments in the thorax of an insect, and bears the first pair of legs. Its principal sclerites (exoskeletal plates) are the pronotum (dorsal), the prosternum (ventral), and the propleuron (lateral) on ea ...
, expanded into often fantastic shapes that enhance their
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the ...
or
mimicry In evolutionary biology, mimicry is an evolved resemblance between an organism and another object, often an organism of another species. Mimicry may evolve between different species, or between individuals of the same species. Often, mimicry f ...
, often resembling plant thorns (thus the commonly used name of "thorn bugs" for a number of treehopper species). The specialised pronotum (or helmet) may not be simply an expansion of the prothoracic sclerite, but a fused pair of dorsal appendages of the first thoracic segment. The treehopper has specialized muscles in its hind femora that unfurl to generate sufficient force to jump. These may be serial homologues of insect wings, which are dorsal appendages of the second and/or third thoracic segments, although this interpretation has been seriously challenged. Evidence for this theory included the development of the helmet, which arises as a pair of appendages attached to each side of the dorsal prothorax by an articulation with muscles and a flexible membrane that allow it to be mobile (also, similar
gene In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a ba ...
s are involved in development of the helmet and the wings). Distinguishing males from females is accomplished only by looking at the genitalia.


Ecology

Treehoppers pierce plant stems with their beaks and feed upon sap. The young can frequently be found on herbaceous shrubs and grasses, while the adults more often frequent hardwood tree species. Excess sap becomes concentrated as honeydew, which often attracts
ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of 22 ...
s. Some species have a well-developed ant mutualism, and these species are normally gregarious as well, which attracts more ants. The ants provide protection from
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
s. Treehoppers mimic thorns to prevent predators from spotting them. Others have formed mutualisms with wasps, such as ''
Parachartergus apicalis ''Parachartergus apicalis'' is a species of wasp in the Polistinae subfamily, found in the Neotropics. It was first described by Johan Christian Fabricius in 1804. In Honduras, they are known as ', which translates into English as "white wings". ...
''. Even
gecko Geckos are small, mostly carnivorous lizards that have a wide distribution, found on every continent except Antarctica. Belonging to the infraorder Gekkota, geckos are found in warm climates throughout the world. They range from . Geckos ar ...
s form mutualistic relations with treehoppers, with whom they communicate by small vibrations of the abdomen. Eggs are laid by the female with her saw-like
ovipositor The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typical ...
in slits cut into the
cambium A cambium (plural cambia or cambiums), in plants, is a tissue layer that provides partially undifferentiated cells for plant growth. It is found in the area between xylem and phloem. A cambium can also be defined as a cellular plant tissue from w ...
or live tissue of stems, though some species lay eggs on top of leaves or stems. The eggs may be
parasitised Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has ...
by
wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. Th ...
s, such as the tiny
fairyflies The Mymaridae, commonly known as fairyflies or fairy wasps, are a family of chalcidoid wasps found in temperate and tropical regions throughout the world. The family contains around 100 genera with 1400 species. Fairyflies are very tiny insects ...
(Mymaridae) and
Trichogrammatidae The Trichogrammatidae are a family of tiny wasps in the Chalcidoidea that include some of the smallest of all insects, with most species having adults less than 1 mm in length, with species of ''Megaphragma'' having an adult body length le ...
. The females of some membracid species sit over their eggs to protect them from predators and parasites, and may buzz their wings at intruders. The females of some gregarious species work together to protect each other's eggs. In at least one species, '' Publilia modesta'', mothers serve to attract ants when
nymph A nymph ( grc, νύμφη, nýmphē, el, script=Latn, nímfi, label=Modern Greek; , ) in ancient Greek folklore is a minor female nature deity. Different from Greek goddesses, nymphs are generally regarded as personifications of nature, are ty ...
s are too small to produce much honeydew. Some other species make feeding slits for the nymphs. Like the adults, the nymphs also feed upon sap, and unlike adults, have an extensible anal tube that appears designed to deposit honeydew away from their bodies. The tube appears to be longer in solitary species rarely attended by ants. It is important for sap-feeding bugs to dispose of honeydew, as otherwise it can become infected with
sooty mould Sooty mold (also spelled sooty mould) is a collective term for different Ascomycete fungi, which includes many genera, commonly ''Cladosporium'' and ''Alternaria''. It grows on plants and their fruit, but also environmental objects, like fences, ...
s. Indeed, one of the evident benefits of ants for '' Publilia concava'' nymphs is that the ants remove the honeydew and reduce such fungal growth. Most species are innocuous to humans, although a few are considered minor pests, such as '' Umbonia crassicornis'' (a thorn bug), the three-cornered alfalfa hopper ('' Spissistilus festinus''), and the buffalo treehopper (''Stictocephala bisonia''), which has been introduced to
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
. The cowbug '' Oxyrachis tarandus'' has been recorded as a pest of ''
Withania somnifera ''Withania somnifera'', known commonly as ashwagandha or winter cherry, is an evergreen shrub in the Solanaceae or nightshade family that grows in India, the Middle East, and parts of Africa. Several other species in the genus ''Withania'' are m ...
'' in India.


Systematics

The diversity of treehoppers has been little researched, and their systematic arrangement is tentative. It seems three main lineages can be distinguished; the Endoiastinae are the most ancient treehoppers, still somewhat resembling
cicada The cicadas () are a superfamily, the Cicadoidea, of insects in the order Hemiptera (true bugs). They are in the suborder Auchenorrhyncha, along with smaller jumping bugs such as leafhoppers and froghoppers. The superfamily is divided into two ...
s. Centrotinae form the second group; they are somewhat more advanced but the pronotum still does not cover the scutellum in almost all of these. The Darninae, Heteronotinae,
Membracinae Membracinae is a subfamily of treehoppers in the family Membracidae. There are more than 40 genera in Membracinae. Genera These 44 genera belong to the subfamily Membracinae: * ''Acanthicoides'' Metcalf, 1952 * ''Aconophora'' Fairmaire, 1846 ...
and
Smiliinae Smiliinae is a subfamily of treehoppers in the family Membracidae. These are Hemiptera, bugs and include about 100 genera in 10 tribes. Tribes and genera These genera belong to the subfamily Smiliinae: * incertae sedis ** ''Antianthe'' Fowler, ...
contain the most
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 treehoppers. Several proposed
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 ...
seem to be
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 ...
. Centronodinae and Nicomiinae might need to be merged into the Centrotinae to result in a
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 ...
group. Treehopper.jpg,
Adults An adult is a human or other animal that has reached full growth. In human context, the term ''adult'' has meanings associated with social and legal concepts. In contrast to a "minor", a legal adult is a person who has attained the age of majori ...
and
nymphs A nymph ( grc, νύμφη, nýmphē, el, script=Latn, nímfi, label=Modern Greek; , ) in ancient Greek folklore is a minor female nature deity. Different from Greek goddesses, nymphs are generally regarded as personifications of nature, are typ ...
of '' Umbonia'' in
Monteverde Monteverde is the twelfth canton of the Puntarenas province of Costa Rica. It is located in the Cordillera de Tilarán mountain range. Roughly a four-hour drive from the Central Valley, Monteverde is one of the country's major ecotourism des ...
,
Costa Rica Costa Rica (, ; ; literally "Rich Coast"), officially the Republic of Costa Rica ( es, República de Costa Rica), is a country in the Central American region of North America, bordered by Nicaragua to the north, the Caribbean Sea to the no ...
Treehopper Horned.JPG, Horned treehopper in
Tumkur Tumkur, officially renamed as Tumakuru, is a city located in the southern part of Indian state of Karnataka. Tumkur is situated at a distance of northwest of Bangalore, the state capital along NH 48 and NH 73. It is the headquarters of the ...
,
India India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the seventh-largest country by area, the second-most populous country, and the most populous democracy in the world. Bounded by the Indian Ocean on the so ...
Treehopper CarpenterAnts.JPG, Treehoppers mating; also some nymphs. Mutualistic carpenter ants present.
Hyderabad Hyderabad ( ; , ) is the capital and largest city of the Indian state of Telangana and the ''de jure'' capital of Andhra Pradesh. It occupies on the Deccan Plateau along the banks of the Musi River (India), Musi River, in the northern part ...
,
India India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the seventh-largest country by area, the second-most populous country, and the most populous democracy in the world. Bounded by the Indian Ocean on the so ...
Treehopper (Membracis bucktoni).jpg, ''
Membracis sanguineoplaga ''Membracis'' is a genus of treehoppers in the family Membracidae. There are more than 50 described species in ''Membracis''. Species These 51 species belong to the genus ''Membracis'': * '' Membracis albolimbata'' Fowler * '' Membracis ambi ...
''
Pantanal,
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...


References


Further reading

*Godoy, C., et al. ''Membrácidos de la América Tropical (Treehoppers of Tropical America''(bilingual, English and Spanish).)''. Santo Domingo de Heredia: INBIO, Inst. Nacional de Biodiversidad. 2006.


External links

* *Dietrich, C. H
Treehopper FAQ.
Section for Biodiversity, Illinois Natural History Survey, Champaign, IL, USA. 2006.
Imagess. Family Membracidae - Treehoppers (United States and Canada).
BugGuide.

*Deitz, L. L. and M. S. Wallace. 2010
Treehoppers: Aetalionidae, Melizoderidae, and Membracidae (Hemiptera).
North Carolina State University Insect Museum. {{Authority control Membracoidea Insect vectors of plant pathogens