''Dactylopius'' is a

genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...

of insect in the superfamily Coccoidea, the

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

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 Juan de Guillebon, better known by his stage name DyE, is a French musician. He is known for the music video of the single "Fantasy Fantasy is a genre of speculative fiction that involves supernatural or Magic (supernatural), magical ele ...

carmine Carmine ()also called cochineal (when it is extracted from the Cochineal, cochineal insect), cochineal extract, crimson Lake pigment, lake, or carmine lake is a pigment of a bright-red color obtained from the aluminium coordination complex, compl ...

. 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, whether pest animals such as insects and mites, weeds, or pathogens affecting animals or plants by using other organisms. It relies on predation, parasitism, herbivory, or ot ...

, and because several are known as

invasive species An invasive species is an introduced species that harms its new environment. Invasive species adversely affect habitats and bioregions, causing ecological, environmental, and/or economic damage. The term can also be used for native spec ...

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. Most species belonging to this genus are covered in thin, sticky strands of wax; ''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 (: cacti, cactuses, or less commonly, cactus) is a member of the plant family Cactaceae (), a family of the order Caryophyllales comprising about 127 genera with some 1,750 known species. The word ''cactus'' derives, through Latin, ...

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

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

'' and '' Grusonia'' species. One species was collected from '' Corynopuntia schottii''. They gather in large clumps mainly in shady, sheltered areas on the cactus pads and feed on the sap. 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, which occurs between each moult (''ecdysis'') until sexual maturity is reached. Arthropods must shed the exoskeleton in order to ...

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 (; seta ; ) are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Depending partly on their form and function, protostome setae may be called macrotrichia, chaetae, ...

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

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 ...

spider Spiders (order (biology), order Araneae) are air-breathing arthropods that have eight limbs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude spider silk, silk. They are the largest order of arachnids and ran ...

s, the larva of the snout moth '' Laetilia coccidivora'', a brown lacewing ('' Symherobius'' sp.), and lady beetles in the genera '' Chilocorus'' 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 considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...

and the others in

North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...

, including

Mexico Mexico, officially the United Mexican States, is a country in North America. It is the northernmost country in Latin America, and borders the United States to the north, and Guatemala and Belize to the southeast; while having maritime boundar ...

and the

southwestern United States The Southwestern United States, also known as the American Southwest or simply the Southwest, is a geographic and cultural list of regions of the United States, region of the United States that includes Arizona and New Mexico, along with adjacen ...

. ''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 Peru, officially the Republic of Peru, is a country in western South America. It is bordered in the north by Ecuador and Colombia, in the east by Brazil, in the southeast by Bolivia, in the south by Chile, and in the south and west by the Pac ...

, 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, also known as the pre-contact era, or as the pre-Cabraline era specifically in Brazil, spans from the initial peopling of the Americas in the Upper Paleolithic to the onset of European col ...

.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 Anti-predator adaptations are mechanisms developed through evolution that assist Predation, prey organisms in their constant struggle against predators. Throughout the animal kingdom, adaptations have evolved for every stage of this struggle, na ...

. It is processed to produce the

pigment A pigment is a powder used to add or alter color or change visual appearance. Pigments are completely or nearly solubility, insoluble and reactivity (chemistry), chemically unreactive in water or another medium; in contrast, dyes are colored sub ...

carmine, which has long been used as a red

food coloring Food coloring, color additive or colorant is any dye, pigment, or substance that imparts color when it is added to food or beverages. Colorants can be supplied as liquids, powders, gels, or pastes. Food coloring is commonly used in commercia ...

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, berry, berries, Bark (botany), bark, leaf, leaves, and wood—and other biological sourc ...

for

textile Textile is an Hyponymy and hypernymy, umbrella term that includes various Fiber, fiber-based materials, including fibers, yarns, Staple (textiles)#Filament fiber, filaments, Thread (yarn), threads, and different types of #Fabric, fabric. ...

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 Domestication is a multi-generational mutualistic relationship in which an animal species, such as humans or leafcutter ants, takes over control and care of another species, such as sheep or fungi, to obtain from them a steady supply of reso ...

and is reared for its product. Cochineals were of value to the pre-Columbian societies of the

Andes The Andes ( ), Andes Mountains or Andean Mountain Range (; ) are the List of longest mountain chains on Earth, longest continental mountain range in the world, forming a continuous highland along the western edge of South America. The range ...

region. Rulers of the

Inca Empire The Inca Empire, officially known as the Realm of the Four Parts (, ), was the largest empire in pre-Columbian America. The administrative, political, and military center of the empire was in the city of Cusco. The History of the Incas, Inca ...

stored supplies of the insects as a

commodity In economics, a commodity is an economic goods, good, usually a resource, that specifically has full or substantial fungibility: that is, the Market (economics), market treats instances of the good as equivalent or nearly so with no regard to w ...

, 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: * the mystique/prestige bestowed upon monarchs ** one or more monarchs, such as kings, queens, emperors, empresses, princes, princesses, etc. *** royal family, the immediate family of a king or queen-regnant, and sometimes h ...

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 The Spanish colonization of the Americas began in 1493 on the Caribbean island of Hispaniola (now Haiti and the Dominican Republic) after the initial 1492 voyage of Genoa, Genoese mariner Christopher Columbus under license from Queen Isabella ...

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 col ...

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

tin Tin is a chemical element; it has symbol Sn () and atomic number 50. A silvery-colored 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, a bar of tin makes a sound, the ...

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

purple Purple is a color similar in appearance to violet light. In the RYB color model historically used in the arts, purple is a secondary color created by combining red and blue pigments. In the CMYK color model used in modern printing, purple is ...

. 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 eco ...

s of

Ecuador Ecuador, officially the Republic of Ecuador, is a country in northwestern South America, bordered by Colombia on the north, Peru on the east and south, and the Pacific Ocean on the west. It also includes the Galápagos Province which contain ...

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 The lemon (''Citrus'' × ''limon'') is a species of small evergreen tree in the ''Citrus'' genus of the flowering plant family Rutaceae. A true lemon is a hybrid of the citron and the bitter orange. Its origins are uncertain, but some ...

juice to turn it scarlet, and the third is rubbed with

wood ash Wood ash is the powder (substance), 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 ...

es to turn it purple.

Biological control

Some cacti are invasive plants, and as

parasites Parasitism is a close relationship between species, where one organism, the parasite, lives (at least some of the time) on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The en ...

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. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...

and

Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...

. The first example of an herbivorous insect ever used for the biological control of a

weed A weed is a plant considered undesirable in a particular situation, growing where it conflicts with human preferences, needs, or goals.Harlan, J. R., & deWet, J. M. (1965). Some thoughts about weeds. ''Economic botany'', ''19''(1), 16-24. Pla ...

was ''D. ceylonicus'', which was released onto ''

Opuntia ficus-indica ''Opuntia ficus-indica'', the Indian fig opuntia, fig opuntia, or prickly pear, is a species of cactus that has long been a domesticated crop plant grown in agricultural economies throughout Arid climate, arid and Semi-arid climate, semiarid part ...

'', then known as ''O. vulgaris'', in

Sri Lanka Sri Lanka, officially the Democratic Socialist Republic of Sri Lanka, also known historically as Ceylon, is an island country in South Asia. It lies in the Indian Ocean, southwest of the Bay of Bengal, separated from the Indian subcontinent, ...

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 language, Spanis ...

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

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 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 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 ...

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

Coccus Bacterial cellular morphologies are the shapes that are characteristic of various types of bacteria and often key to their identification. Their direct examination under a light microscope enables the classification of these bacteria (and archaea ...

'' in 2001 and ''D. gracilipilus'' was newly described in 2012. Species include: *'' Dactylopius austrinus'' *'' Dactylopius bassi'' *'' Dactylopius ceylonicus'' *'' Dactylopius coccus'' – true cochineal *'' Dactylopius confertus'' *'' Dactylopius confusus'' *'' Dactylopius gracilipilus'' *'' Dactylopius opuntiae'' *'' Dactylopius salmianus'' *'' Dactylopius tomentosus'' *'' Dactylopius zimmermanni''