''Dactylopius'' is a

genus 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 nom ...

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.) (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 sessile parasite native to tropical and subtropical South America through North Americ ...

(''Dactylopius coccus''). The cochineal is an insect of economic and historical importance as a main source of the red dye carmine. 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, and because several are known as invasive species.

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 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 giv ...

; ''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 Gree ...

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

. 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 ...

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. ...

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 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 ...

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

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

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 relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the sout ...

and the others in North America, 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. ''D. coccus'' apparently has a disjunct distribution 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 = National seal , national_motto = "Firm and Happy f ...

, but not in the region between. Molecular phylogenetic 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 original settlement of North and South America in the Upper Paleolithic period through European colonization, which began with Christopher Columbus's voyage of 1492. Usually, ...

.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 prey organisms in their constant struggle against predators. Throughout the animal kingdom, adaptations have evolved for every stage of this struggle, namely by avo ...

. 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 compou ...

carmine, which has long been used as a red

food coloring Food coloring, or color additive, is any dye, pigment, or substance that imparts color when it is added to food or drink. They come in many forms consisting of liquids, powders, gels, and pastes. Food coloring is used in both commercial food ...

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. Archa ...

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 Domestication is a sustained multi-generational relationship in which humans assume a significant degree of control over the reproduction and care of another group of organisms to secure a more predictable supply of resources from that group. A ...

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 adm ...

stored supplies of the insects as a commodity, 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 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, 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, ...

. 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: ' ...

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 trees in the flowering plant family Rutaceae, native to Asia, primarily Northeast India (Assam), Northern Myanmar or China. The tree's ellipsoidal yellow fruit is used for culin ...

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 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 ha ...

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. 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, "a plant in the wrong place", or a plant growing where it is not wanted.Harlan, J. R., & deWet, J. M. (1965). Some thoughts about weeds. ''Economic botany'', ''19''(1), 16-24. ...

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 and semiarid parts of the world. ''O. ficus-indica ...

'', then known as ''O. vulgaris'', in Sri Lanka 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 des ...

'', 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 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. 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 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 ( ...

'' – true cochineal *'' Dactylopius confertus'' *'' Dactylopius confusus'' *'' Dactylopius gracilipilus'' *'' Dactylopius opuntiae'' *'' Dactylopius salmianus'' *'' Dactylopius tomentosus'' *'' Dactylopius zimmermanni''