Dactylopius Confusus
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''Dactylopius'' is a genus 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 (''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; ''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 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. 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 ( instars), 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 setae 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, spiders, 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 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 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 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, 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.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 carmine, which has long been used as a red food coloring and a natural dye for textiles. ''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 region. Rulers of the Inca Empire 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 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 salts or acids produce a bright
scarlet Scarlet may refer to: * Scarlet (cloth), a type of woollen cloth common in medieval England * Scarlet (color), a bright tone of red that is slightly toward orange, named after the cloth * Scarlet (dye), the dye used to give the cloth its color * ...
, while iron or alkaline compounds turn it a deep purple. The Salasacas of Ecuador 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 ashes 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 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'', 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 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'' in 2001 and ''D. gracilipilus'' was newly described in 2012. Species include: *'' Dactylopius austrinus'' *''
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'' 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 coccus'' – true cochineal *''
Dactylopius confertus ''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 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 opuntiae'', also known as the prickly pear cochineal, is a species of scale insect in the family Dactylopiidae. Taxonomy ''Dactylopius opuntiae'' was first identified by Cockerell as ''Coccus cacti opuntiae'' after he collected it ...
'' *''
Dactylopius salmianus ''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 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.) ...
'' *''
Dactylopius zimmermanni ''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.) ...
''


References


Further reading

*Chávez Moreno, C. K
The ''Opuntia'' (Cactaceae) and ''Dactylopius'' (Hemiptera: Dactylopiidae) in Mexico: a historical perspective of use, interaction and distribution with particular emphasis on chemical and phylogenetic aspects of the ''Dactylopius'' species.
VII International Congress on Cactus & Cochineal, Agadir, Morocco, October 22, 2010. {{Taxonbar, from=Q677616 Hemiptera of South America Dactylopiidae Sternorrhyncha genera Taxa named by Achille Costa