''Dactylopius'' is a

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...

of insect in the superfamily Coccoidea, the

scale insect Scale insects are small insects of the order Hemiptera, suborder Sternorrhyncha. Of dramatically variable appearance and extreme sexual dimorphism, they comprise the infraorder Coccomorpha which is considered a more convenient grouping than the ...

s. It is the only genus in the family Dactylopiidae.Van Dam, A. R. and B. May. (2012)
A new species of ''Dactylopius'' Costa (''Dactylopius gracilipilus'' sp. nov.) (Hemiptera: Coccoidea: Dactylopiidae) from the Chihuahuan Desert, Texas, U.S.A.
''Zootaxa'' 3573: 33–39. These insects are known commonly as cochineals,Ramírez-Puebla, S. T., et al. (2010)
Molecular phylogeny of the genus ''Dactylopius'' (Hemiptera: Dactylopiidae) and identification of the symbiotic bacteria.
''Environmental Entomology'' 39(4), 1178-83.Chávez-Moreno, C. K., et al. (2011)
Distribution and habitat in Mexico of ''Dactylopius'' Costa (Hemiptera: Dactylopiidae) and their cacti hosts (Cactaceae: Opuntioideae).
''Neotropical Entomology'' 40(1), 62-71. a name that also specifically refers to the best-known species, the

cochineal The cochineal ( , ; ''Dactylopius coccus'') is a scale insect in the suborder Sternorrhyncha, from which the natural dye carmine is derived. A primarily sessility (motility), sessile parasitism, parasite native to tropical and subtropical Sout ...

(''Dactylopius coccus''). The cochineal is an insect of economic and historical importance as a main source of the red

dye A dye is a colored substance that chemically bonds to the substrate to which it is being applied. This distinguishes dyes from pigments which do not chemically bind to the material they color. Dye is generally applied in an aqueous solution an ...

carmine Carmine ()also called cochineal (when it is extracted from the cochineal insect), cochineal extract, crimson lake, or carmine lake is a pigment of a bright-red color obtained from the aluminium complex derived from carminic acid. Specific code n ...

. It has reportedly been used for this purpose in the Americas since the 10th century. Genus ''Dactylopius'' is also important because several species have been used as agents of

biological pest control Biological control or biocontrol is a method of controlling pests, such as insects, mites, weeds, and plant diseases, using other organisms. It relies on predation, parasitism, herbivory, or other natural mechanisms, but typically also invo ...

, and because several are known as

invasive species An invasive species otherwise known as an alien is an introduced organism that becomes overpopulated and harms its new environment. Although most introduced species are neutral or beneficial with respect to other species, invasive species ad ...

Description

In general, cochineals are waxy gray scaled insects, with the key characteristic that adult females release the bright red substance when crushedFamily:Dactylopiidae.
Scale Insects. Systematic Entomology Laboratory, USDA, ARS. known as

carminic acid Carminic acid (C22H20O13) is a red glucosidal hydroxyanthrapurin that occurs naturally in some scale insects, such as the cochineal, Armenian cochineal, and Polish cochineal. The insects produce the acid as a deterrent to predators. An aluminu ...

. Most species belonging to this genus are covered in thin, sticky strands of

wax Waxes are a diverse class of organic compounds that are lipophilic, malleable solids near ambient temperatures. They include higher alkanes and lipids, typically with melting points above about 40 °C (104 °F), melting to give low ...

; ''D. coccus'' is one such species which lacks these traits. Common with the characteristics of most scaled insects, the species within this genus too show sexual dimorphism between male and female individuals. Where females of the genus possess a simple morphology (standard body, with a head, mouth, legs, etc.), though lacking wings of any kind; the males of the genus are the opposite, possessing a singular set of wings, and no mouth at all. (See '' D. coccus'' for example.)

Biology

Cochineals live on

cactus A cactus (, or less commonly, cactus) is a member of the plant family Cactaceae, a family comprising about 127 genera with some 1750 known species of the order Caryophyllales. The word ''cactus'' derives, through Latin, from the Ancient Greek ...

plants, especially prickly pears (''Opuntia'' spp.), as well as ''

Cylindropuntia ''Cylindropuntia'' is a genus of cacti (family Cactaceae), containing species commonly known as chollas, native to northern Mexico and the Southwestern United States. They are known for their barbed spines that tenaciously attach to skin, fur, a ...

'' and ''

Grusonia ''Grusonia'' is a genus of opuntioid cacti (family Cactaceae), originating from the North American Deserts in Southwest United States and northern Mexico, including Baja California. Authors differ on precise boundaries of the genus, which has b ...

'' species. One species was collected from ''

Corynopuntia schottii ''Grusonia'' is a genus of opuntioid cacti (family Cactaceae), originating from the North American Deserts in Southwest United States and northern Mexico, including Baja California. Authors differ on precise boundaries of the genus, which has be ...

''. They gather in large clumps mainly in shady, sheltered areas on the cactus pads and feed on the

sap Sap is a fluid transported in xylem cells (vessel elements or tracheids) or phloem sieve tube elements of a plant. These cells transport water and nutrients throughout the plant. Sap is distinct from latex, resin, or cell sap; it is a separa ...

. Some species also infest the cactus fruits, flowers, stems, and roots; they are often found on the roots during times of the year when they are less active. Females have three developmental stages (

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

s), and males have five. They reproduce continuously year-round, with up to six generations per year. Eggs hatch very soon after they are laid, sometimes within minutes, and probably even occasionally while they are still inside the female. ''D. tomentosus'' carries its eggs on its body in a mesh of fibers. The wax filaments are produced from

seta In biology, setae (singular seta ; from the Latin word for "bristle") are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Annelid setae are stiff bristles present on the body. Th ...

e on the juvenile insect's head. These webby filaments likely aid in dispersal of the insects by catching the wind.Mow, V., et al. (1982)
Wind dispersal and settling of first-instar crawlers of the cochineal insect ''Dactylopius austrinus'' (Homoptera: Coccoidea: Dactylopiidae).
''Ecological Entomology'' 7(4), 409-19. ''Dactylopius'' species can be found on cacti in many types of dry habitat, including forests, grasslands, cultivated fields, disturbed and weedy areas, and gardens. They share cactus host plants with various ants,

spider Spiders ( order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species ...

s, the larva of the snout moth ''

Laetilia coccidivora ''Laetilia coccidivora'', the scale-feeding snout moth, is a species of snout moth The Pyralidae, commonly called pyralid moths, snout moths or grass moths, are a family of Lepidoptera in the ditrysian superfamily Pyraloidea. In many (particul ...

'', a brown lacewing ('' Symherobius'' sp.), and

lady beetles Coccinellidae () is a widespread family (biology), family of small beetles ranging in size from . They are commonly known as ladybugs in North America and ladybirds in Great Britain. Some Entomology, entomologists prefer the names ladybird be ...

in the genera ''

Chilocorus ''Chilocorus'' is a genus of beetles belonging to the family Coccinellidae, subfamily Chilocorinae. List of species These species belong to the genus ''Chilocorus''. * '' Chilocorus bipustulatus'' (Linnaeus, 1758) - Heather ladybird * '' ...

'' and '' Hyperaspis''.

Distribution

The genus is native to the Americas, with some originating in

South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...

and the others in

North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...

, including

Mexico Mexico (Spanish: México), officially the United Mexican States, is a country in the southern portion of North America. It is bordered to the north by the United States; to the south and west by the Pacific Ocean; to the southeast by Guatema ...

and the

southwestern United States The Southwestern United States, also known as the American Southwest or simply the Southwest, is a geographic and cultural region of the United States that generally includes Arizona, New Mexico, and adjacent portions of California, Colorado, Ne ...

. ''D. coccus'' apparently has a

disjunct distribution In biology, a taxon with a disjunct distribution is one that has two or more groups that are related but considerably separated from each other geographically. The causes are varied and might demonstrate either the expansion or contraction of a s ...

today, occurring in Mexico and also in

Peru , image_flag = Flag of Peru.svg , image_coat = Escudo nacional del Perú.svg , other_symbol = Great Seal of the State , other_symbol_type = Seal (emblem), National seal , national_motto = "Fi ...

, but not in the region between.

Molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...

studies of the genus suggest that ''D. coccus'' originated in South America and was introduced to Mexico with various agricultural products during the

Pre-Columbian era In the history of the Americas, the pre-Columbian era spans from the Migration to the New World, original settlement of North and South America in the Upper Paleolithic period through European colonization of the Americas, European colonization, w ...

.Rodríguez, L. C., et al. (2001)
Direction of dispersion of cochineal (''Dactylopius coccus'' Costa) within the Americas.
''Antiquity'' 75, 73-77. The genus is now distributed throughout much of the world due to accidental and intentional introductions.

Cochineal dye

The insects produce carminic acid as an antipredator adaptation. It is processed to produce the

pigment A pigment is a colored material that is completely or nearly insoluble in water. In contrast, dyes are typically soluble, at least at some stage in their use. Generally dyes are often organic compounds whereas pigments are often inorganic compo ...

carmine, which has long been used as a red food coloring and a

natural dye Natural dyes are dyes or colorants derived from plants, invertebrates, or minerals. The majority of natural dyes are vegetable dyes from plant sources—roots, berries, bark, leaves, and wood—and other biological sources such as fungi. Archaeol ...

for

textile Textile is an umbrella term that includes various fiber-based materials, including fibers, yarns, filaments, threads, different fabric types, etc. At first, the word "textiles" only referred to woven fabrics. However, weaving is not the ...

s. ''Dactylopius coccus'', the true cochineal, is the species most commonly used today and historically, because it has a higher carminic acid content and yields a better quality pigment than its congeners. The insect has been domesticated and is reared for its product. Cochineals were of value to the pre-Columbian societies of the

Andes The Andes, Andes Mountains or Andean Mountains (; ) are the longest continental mountain range in the world, forming a continuous highland along the western edge of South America. The range is long, wide (widest between 18°S – 20°S ...

region. Rulers of the

Inca Empire The Inca Empire (also known as the Incan Empire and the Inka Empire), called ''Tawantinsuyu'' by its subjects, (Quechua for the "Realm of the Four Parts", "four parts together" ) was the largest empire in pre-Columbian America. The admin ...

stored supplies of the insects as a

commodity In economics, a commodity is an economic good, usually a resource, that has full or substantial fungibility: that is, the market treats instances of the good as equivalent or nearly so with no regard to who produced them. The price of a comm ...

, as they were the only source of red dye for clothing and other textiles.Roquero, A
Identification of red dyes in textiles from the Andean Region.
Paper 129 In: Proceedings, Textile Society of America 11th Biennial Symposium, Lincoln, Nebraska, October 6–9, 2010. Red was the color of

royalty Royalty may refer to: * Any individual monarch, such as a king, queen, emperor, empress, etc. * Royal family, the immediate family of a king or queen regnant, and sometimes his or her extended family * Royalty payment for use of such things as int ...

among the Incas.Phipps, E. and N. Shibayama
Tracing cochineal through the collection of the Metropolitan Museum.
Paper 44 In: Proceedings, Textile Society of America 12th Biennial Symposium, Honolulu, September 4–7, 2008. After

Spanish colonization of the Americas Spain began colonizing the Americas under the Crown of Castile and was spearheaded by the Spanish . The Americas were invaded and incorporated into the Spanish Empire, with the exception of Brazil, British America, and some small regions ...

began, cochineals were shipped worldwide as a commercial product. The dried bodies of the female insects are roughly 12 to 16% carminic acid.Reyes-Salas, O., et al. (2011)
Titrimetric and polarographic determination of carminic acid and its quantification in cochineal (''Dactylopius coccus'') extracts.
''Journal of the Mexican Chemical Society'' 55(2), 88-93. The carmine obtained is a shade of

crimson Crimson is a rich, deep red color, inclining to purple. It originally meant the color of the kermes dye produced from a scale insect, ''Kermes vermilio'', but the name is now sometimes also used as a generic term for slightly bluish-red colo ...

, and this can be turned different shades by adding metal ions or changing the pH;

tin Tin is a chemical element with the symbol Sn (from la, stannum) and atomic number 50. Tin is a silvery-coloured metal. Tin is soft enough to be cut with little force and a bar of tin can be bent by hand with little effort. When bent, t ...

salts or acids produce a bright scarlet, while iron or alkaline compounds turn it a deep

purple Purple is any of a variety of colors with hue between red and blue. In the RGB color model used in computer and television screens, purples are produced by mixing red and blue light. In the RYB color model historically used by painters, pu ...

. The

Salasaca Salasaca is a community and an indigenous people located in the Tungurahua Province in the center of Ecuador, halfway along the road from Ambato to Baños. The Salasaca speak Spanish and their traditional language of Quichua. Their main econom ...

s of

Ecuador Ecuador ( ; ; Quechua: ''Ikwayur''; Shuar: ''Ecuador'' or ''Ekuatur''), officially the Republic of Ecuador ( es, República del Ecuador, which literally translates as "Republic of the Equator"; Quechua: ''Ikwadur Ripuwlika''; Shuar: ''Eku ...

still use this process to color their textiles, particularly a red woolen garment worn by women around the shoulders, which each woman personalizes by choosing a different shade of red. They collect cochineals, probably ''D. confusus'', from the wild, press them into cakes, and use the dried cakes to dye the garments three at a time. One is left crimson, one is soaked in lemon juice to turn it scarlet, and the third is rubbed with

wood ash Wood ash is the powdery residue remaining after the combustion of wood, such as burning wood in a fireplace, bonfire, or an industrial power plant. It is largely composed of calcium compounds along with other non-combustible trace elements presen ...

es to turn it purple.

Biological control

Some cacti are invasive plants, and as parasites of cacti, ''Dactylopius'' species have been employed as agents of biological pest control in

Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...

and

Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...

. The first example of an herbivorous insect ever used for the biological control of a weed was ''D. ceylonicus'', which was released onto '' Opuntia ficus-indica'', then known as ''O. vulgaris'', in

Sri Lanka Sri Lanka (, ; si, ශ්‍රී ලංකා, Śrī Laṅkā, translit-std=ISO (); ta, இலங்கை, Ilaṅkai, translit-std=ISO ()), formerly known as Ceylon and officially the Democratic Socialist Republic of Sri Lanka, is an ...

in 1863.Volchansky, C. R., et al. (1999)
Host-plant affinities of two biotypes of ''Dactylopius opuntiae'' (Homoptera: Dactylopiidae): enhanced prospects for biological control of ''Opuntia stricta'' (Cactaceae) in South Africa.
''Journal of Applied Ecology'' 36(1), 85-91. ''D. opuntiae'' has been effective against the weedy prickly pear ''

Opuntia stricta ''Opuntia stricta'' is a species of large cactus that is endemic to the subtropical and tropical coastal areas of the Americas, especially around the Caribbean. Common names include erect prickly pear and nopal estricto (Spanish). The first desc ...

'', and ''D. austrinus'' is used to control ''

Opuntia aurantiaca ''Opuntia aurantiaca'', commonly known as tiger-pear, jointed cactus or jointed prickly-pear, is a species of cactus from South America. The species occurs naturally in Argentina, Paraguay and Uruguay and is considered an invasive species in Afr ...

''.

As a pest

''O. ficus-indica'' is a cactus cultivated as a food crop, and it is the most commonly noted host plant of ''Dactylopius'' species. ''D. coccus'' can be found in fields of the cultivated plant and on specimens in the wild. ''D. opuntiae'' is an economically important

pest Pest or The Pest may refer to: Science and medicine * Pest (organism), an animal or plant deemed to be detrimental to humans or human concerns ** Weed, a plant considered undesirable * Infectious disease, an illness resulting from an infection ** ...

of this cactus in many regions. Vanegas-Rico, J. M., et al. (2010)
Enemigos naturales de ''Dactylopius opuntiae'' (Cockerell) en ''Opuntia ficus-indica'' (L.) Miller en el centro de México.
''Acta Zoológica Mexicana'' 26(2), 415-33.

Diversity

This genus is

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 ...

. There are eleven species. In addition to the nine that long composed the genus, ''D. bassi'' was transferred from genus ''

Coccus A coccus (plural cocci) is any bacterium or archaeon that has a spherical, ovoid, or generally round shape. Bacteria are categorized based on their shapes into three classes: cocci (spherical-shaped), bacillus (rod-shaped) and spiral ( of whi ...

'' in 2001 and ''D. gracilipilus'' was newly described in 2012. Species include: *''

Dactylopius austrinus ''Dactylopius'' is a genus of insect in the superfamily Coccoidea, the scale insects. It is the only genus in the family Dactylopiidae.Van Dam, A. R. and B. May. (2012)A new species of ''Dactylopius'' Costa (''Dactylopius gracilipilus'' sp. nov.) ...

'' *''

Dactylopius bassi ''Dactylopius'' is a genus of insect in the superfamily Coccoidea, the scale insects. It is the only genus in the family Dactylopiidae.Van Dam, A. R. and B. May. (2012)A new species of ''Dactylopius'' Costa (''Dactylopius gracilipilus'' sp. nov.) ...

'' *'' Dactylopius ceylonicus'' *''

Dactylopius coccus The cochineal ( , ; ''Dactylopius coccus'') is a scale insect in the suborder Sternorrhyncha, from which the natural dye carmine is derived. A primarily sessile parasite native to tropical and subtropical South America through North America (M ...

'' – true cochineal *'' Dactylopius confertus'' *'' Dactylopius confusus'' *''

Dactylopius gracilipilus ''Dactylopius'' is a genus of insect in the superfamily Coccoidea, the scale insects. It is the only genus in the family Dactylopiidae.Van Dam, A. R. and B. May. (2012)A new species of ''Dactylopius'' Costa (''Dactylopius gracilipilus'' sp. nov.) ...

'' *'' Dactylopius opuntiae'' *'' Dactylopius salmianus'' *''

Dactylopius tomentosus ''Dactylopius'' is a genus of insect in the superfamily Coccoidea, the scale insects. It is the only genus in the family Dactylopiidae.Van Dam, A. R. and B. May. (2012)A new species of ''Dactylopius'' Costa (''Dactylopius gracilipilus'' sp. nov.) ...