Treehoppers (more precisely typical treehoppers to distinguish them from the Aetalionidae) and thorn bugs are members of the

family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...

Membracidae, a group of

insect Insects (from Latin ') are Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (Insect morphology#Head, head, ...

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 Leafhopper is the common name for any species from the family (biology), family Cicadellidae: based on the type genus ''Cicadella''. These minute insects, colloquially known as hoppers, are plant feeders that suck plant sap from grass, shrubs, or ...

s. About 3,200

species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...

of treehoppers in over 400

genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...

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 continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...

. Individual treehoppers usually live for only a few months.

Morphology

Treehoppers, due to their unusual appearance, have long interested

naturalist Natural history is a domain of inquiry involving organisms, including animals, fungi, and plants, in their natural environment, leaning more towards observational than experimental methods of study. A person who studies natural history is cal ...

s. They are best known for their enlarged and ornate pronotum, 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 b ...

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. In the simples ...

, often resembling plant thorns (thus the commonly used name of "thorn bugs" for a number of treehopper species). Treehoppers have specialized muscles in the hind femora that unfurl to generate sufficient force to jump. It had been suggested that the pronotal "helmet" could be serial homologues of insect wings, but this interpretation has been refuted by several later studies, such as Mikó et al. (2012) and Yoshizawa (2012). Treehopper nymphs can be recognised by the tube-like ninth abdominal segment, through which the tenth and eleventh segments can be exerted in defence or to provide honeydew to other animals (explained further in the next section). The tube is longer (compared to the rest of the body) in early

instar An instar (, from the Latin '' īnstar'' 'form, likeness') is a developmental stage of arthropods, such as insects, which occurs between each moult (''ecdysis'') until sexual maturity is reached. Arthropods must shed the exoskeleton in order to ...

s compared to late instars.

Ecology

Treehoppers have pointy, tube-shaped mouthparts that they use to pierce plant stems 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 Eusociality, eusocial insects of the Family (biology), family Formicidae and, along with the related wasps and bees, belong to the Taxonomy (biology), order Hymenoptera. Ants evolved from Vespoidea, vespoid wasp ancestors in the Cre ...

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 in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...

s. Treehoppers mimic thorns to prevent predators from spotting them. Others have formed mutualisms with wasps, such as '' Parachartergus apicalis''. 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. They range from . Geckos are unique among lizards ...

s form mutualistic relations with treehoppers, with whom they communicate by small vibrations of the abdomen. Mutualisms are not done only for protection against predators. Nymphs of the treehopper '' Publilia concava'' have higher survivorship in the presence of ants even when predators are absent. This is suspected to be because uncollected honeydew leads to the growth of sooty mould, which may hinder excretion by treehoppers and

photosynthesis Photosynthesis ( ) is a system of biological processes by which photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical energy necessary to fuel their metabo ...

by their host plants. Ant collection of honeydew thus allows treehoppers to feed more (the feeding facilitation hypothesis). 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 typica ...

in slits cut into the

cambium A cambium (: cambiums or cambia), 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 whic ...

or live tissue of stems, though some species lay eggs on top of leaves or stems. The eggs may be parasitised 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 ...

s, such as the tiny fairyflies (Mymaridae) and Trichogrammatidae. 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 (; ; sometimes spelled nymphe) is a minor female nature deity in ancient Greek folklore. Distinct from other Greek goddesses, nymphs are generally regarded as personifications of nature; they are typically tied to a specific place, land ...

s are too small to produce much honeydew. Some other species make feeding slits for the nymphs. 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

. The cowbug '' Oxyrachis tarandus'' has been recorded as a pest of ''

Withania somnifera ''Withania somnifera'', known commonly as ashwagandha, is an evergreen shrub in the Solanaceae family that is native to the Middle East and North Africa, other African regions, southern Europe, and Indian subcontinent. Several other species in ...

'' 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

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 and Smiliinae 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 ha ...

ic treehoppers. Several proposed subfamilies seem to 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 ...

. Centronodinae and Nicomiinae might need to be merged into the Centrotinae to result in a

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. Treehopper.jpg, Adults and

nymphs A nymph (; ; sometimes spelled nymphe) is a minor female nature deity in ancient Greek folklore. Distinct from other Greek goddesses, nymphs are generally regarded as personifications of nature; they are typically tied to a specific place, land ...

of '' Umbonia'' in

Monteverde Monteverde is the twelfth Cantons of Costa Rica, canton of the Puntarenas Province, Puntarenas Provinces of Costa Rica, province of Costa Rica, located in the Cordillera de Tilarán (Tilarán range). Roughly a four-hour drive from the Costa Rica ...

Costa Rica Costa Rica, officially the Republic of Costa Rica, is a country in Central America. It borders Nicaragua to the north, the Caribbean Sea to the northeast, Panama to the southeast, and the Pacific Ocean to the southwest, as well as Maritime bo ...

Treehopper Horned.JPG, Horned treehopper in

Tumkur Tumkur, officially Tumakuru, is a city and headquarters of Tumakuru district in the Karnataka state of India. Tumkur is known for Siddaganga Matha. Tumkur hosts India's first mega food park, a project of the ministry of food processing. The Ind ...

India India, officially the Republic of India, is a country in South Asia. It is the List of countries and dependencies by area, seventh-largest country by area; the List of countries by population (United Nations), most populous country since ...

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. It occupies on the Deccan Plateau along the banks of the Musi River (India), Musi River, in the northern part of Southern India. With an average altitude of , much ...

Treehopper (Membracis bucktoni).jpg, '' Membracis sanguineoplaga''
Pantanal,

Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is the world's List of countries and dependencies by area, fifth-largest country by area and the List of countries and dependencies by population ...

References

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 Insect vectors of plant pathogens

Morphology

Ecology

Systematics

References

Further reading

External links