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Family Dactylopiidae

Dactylopius Costa

NOMENCLATURE:

Diaprosteci Costa, 1828: 7. Unavailable name.

Dactylopius Costa, 1829: 16. Type species: Dactylopius coccus Costa. Subsequently designated by ICZN, 1983: 77. Notes: Opinion 1247 of the International Commission on Zoological Nomenclature (ICZN, 1983) set aside previous designations of type species for Dactylopius and designated Dactylopius coccus Costa, 1829 as the type-species. The above Opinion placed Dactylopius Costa, 1829 on the Official List of Generic Names in Zoology, and Dactylopius coccus Costa, 1829 on the Official List of Specific Names in Zoology.

Diaprostocetus; Westwood, 1840: 447. Misspelling of genus name.

Diaprostecha; Targioni Tozzetti, 1868: 697. Misspelling of genus name.

Diaprostechus; Targioni Tozzetti, 1868: 725. Misspelling of genus name.

Diasprostetus; Signoret, 1868: 842. Misspelling of genus name.

Diasprotecus; Signoret, 1868: 511. Misspelling of genus name.

Dactylopus; Signoret, 1875c: 499. Misspelling of genus name.

Diasprostethus; Signoret, 1877: 658. Misspelling of genus name.

Diaprostecie; Cockerell, 1902: 454. Misspelling of genus name.

Diaprostecus; Waterhouse, 1902: 410. Misspelling of genus name.

BIOLOGY: All described species of this genus are known to develop only on host plants of the Cactaceae.

GENERAL REMARKS: Definition and characters by Ferris (1955a), De Lotto (1974a), Perez Guerra & Kosztarab (1992) and by Gill (1993).

SYSTEMATICS: This genus currently accommodates species of cochineal scales, family Dactylopiidae. However, during the latter decades of 19th century and early 20's, many species, which belong to the mealybug family, Pseudococcidae, have been described in this genus. The history, taxonomy and nomenclature of this confusion has been extensively discussed by Ferris (1955a), Morrison & Morrison (1966), De Lotto (1974a), Miller (1974), Danzig & Kerzhner (1981), and Opinion 1247 (ICZN, 1983).

ECONOMIC IMPORTANCE AND CONTROL: Chavez-Moreno et al. (2010) studied five species of Dactylopius collected in thirteen states of Mexico and two provinces of Argentina and analyzed using high-performance liquid chromatography with a photodiode array detector. This analysis allowed each species to be identified on the basis of differences in their metabolic profiles. Cluster analysis and principal component analysis differentiated the species by localities and host plant. These two multivariate data analysis techniques were complementary and confirmed the grouping of all analyzed Dactylopius samples. For all species, carminic acid, identified by reference to a commercial sample, was the major compound present in significant amounts, making all five species potential sources of colorant. In addition, each species could be differentiated by the presence of other minor compounds.

KEYS: De Haro & Claps 1995: 16-17 (female) [world]; Gill 1993: 148 (female) [USA, California]; Perez Guerra & Kosztarab 1992: 35-36 (female) [Species of Dactylopius]; De Lotto 1974a (female) [world]; Ferris 1955a: 86 (female) [North America].

CITATIONS: Ashmea1891 [taxonomy: 95]; Atkins1886 [taxonomy: 285,288]; BenDov2006b [taxonomy, catalogue: 211]; ChavezTeCa2011 [distribution: 64]; ChavezTeFr2010 [chemistry, carminic acid: 671-679]; Cocker1893dd [taxonomy: 1041-1049]; Cocker1902k [taxonomy: 453-454]; Comsto1881a [taxonomy: 345]; Costa1829 [taxonomy: 16]; DanzigKe1981 [taxonomy: 79-83]; DeHaroCl1995 [taxonomy, description, host plants, economic importance: 1-19]; DeLott1974a [taxonomy, description: 167-170]; Fernal1903b [catalogue: 80-82]; Ferris1919a [taxonomy: 19]; Ferris1937 [taxonomy: 5]; Ferris1955a [taxonomy, description: 85-86]; Ferris1957c [taxonomy: 85]; Gill1993 [taxonomy, description: 146-148]; Green1896e [taxonomy: 17]; Green1912a [taxonomy, description: 79-93]; Green1922 [taxonomy: 357]; GullanCo2001 [taxonomy: 91-99]; Howell1976 [taxonomy: 188]; ICZN1983 [taxonomy: 77-80]; KozarDr1998f [catalogue: 169]; MacGreSa1984 [taxonomy: 217]; Mamet1949 [taxonomy: 7]; Mann1969 [host, distribution, life history: 1-158]; Maskel1887a [taxonomy: 89]; Maskel1894b [taxonomy: 86]; Miller1974b [taxonomy: 146-153]; MorrisMo1966 [taxonomy: 54-55]; PerezGKo1992 [taxonomy, description: 1-87]; PortilVi2006 [taxonomy, host, distribution: 249-255]; Signor1875c [taxonomy: 339-340]; SimmonBe1966 [distribution, biological control: 183-189]; VanDamMa2012 [description, distribution, economic importance, taxonomy: 33,35]; Willia1969a [taxonomy: 324].

Dactylopius austrinus De Lotto

NOMENCLATURE:

Dactylopius austrinus De Lotto, 1974a: 174. Type data: SOUTH AFRICA: Cape Province, on Opuntia aurantiaca. Holotype female. Type depository: Pretoria: South African National Collection of Insects, South Africa. Described: female. Illust.

HOSTS: Cactaceae: Opuntia anacantha anacantha [ClapsDe2001], Opuntia anacantha kiska-loro, Opuntia aurantiaca [DeLott1974a, MoranCo1979, Hoskin1984, PerezGKo1992], Opuntia canina [PerezGKo1992], Opuntia kiska-loro [PerezGKo1992], Opuntia pampeana [PerezGKo1992], Opuntia retrorsa [PerezGKo1992], Opuntia sulphurea [ClapsDe2001].

DISTRIBUTION: Afrotropical: South Africa [DeLott1974a, MoranCo1979, MoranGuWa1982, PerezGKo1992]. Australasian: Australia (New South Wales [DeLott1974a, Hoskin1984, PerezGKo1992]). Neotropical: Argentina (Catamarca [PerezGKo1992], Jujuy [PerezGKo1992], La Rioja [PerezGKo1992], Mendoza [DeLott1974a]).

BIOLOGY: Life history of the male and female in South Africa studied and described by Moran & Cobby (1979). Hosking (1984) studied the effect of temperature on the population growth potential of this scale, on Opuntia aurantiaca in Australia.

GENERAL REMARKS: Description and illustration of adult female by Perez Guerra & Kosztarab (1992). Portillo (1995) distinguished between the fine cochineal species Dactylopius coccus and a group of wild cochineal species comprising D. austrinus, D. ceylonicus, D. confertus, D. confusus, D. opuntiae, D. salmianus, D. tomentosus and D. zimmermani.

STRUCTURE: Gunn (1978) described for the first time sexual dimorphism in crawlers of this species, and in the Dactylopiidae.

ECONOMIC IMPORTANCE AND CONTROL: Imported for the biological control of jointed cactus, Opuntia aurantiaca in South Africa (Moran & Cobby, 1979), where it is regarded the primary biocontrol agent (Moran & Zimmermann, 1991). Introduced into Australia for the biological control of tiger pear, Opuntia aurantiaca (Hosking, 1984).

KEYS: De Haro & Claps 1995: 16-17 (female) [world]; Perez Guerra & Kosztarab 1992: 35-36 (female) [Species of Dactylopius]; De Lotto 1974a: 173-174 (female) [world].

CITATIONS: BenDov2006b [taxonomy, catalogue: 212]; ClapsDe2001 [host, distribution: 79]; ControJoCa1981 [host, distribution, biological control: 1]; CookGu2004 [taxonomy: 444]; Cullen1995 [biological control: 103-109]; DeHaroCl1995 [taxonomy: 16-17]; DeLott1974a [taxonomy, description, illustration, host, distribution: 174, 175]; GullanCo2001 [taxonomy: 91-99]; GullanCo2007 [taxonomy: 413-425]; Gunn1978 [taxonomy, morphology: 333-338]; GwiazdVaDe2006 [phylogenetics: 16]; Hoskin1984 [life history, biological control, economic importance, host, distribution: 133-139]; KlugeZiCi1986 [biological control: 295-303]; Koszta1987 [economic importance, biological control: 218]; MoranCo1979 [life history, biological control, host, distribution: 629-636]; MoranGuWa1982 [life history, biological control, host, distribution: 409-419]; MoranNeHo1986 [biological control: 261-268]; MoranZi1991 [host, distribution, economic importance, biological control: 5-27]; NanDeWu2013 [phylogenetics: 173]; PerezGKo1992 [taxonomy, description, illustration, host, distribution: 36-40]; Portil1995 [host, distribution: 62-65]; ZimmerBuAn1974 [host, distribution, biological control, economic importance: 204-209]; ZimmerMo1982 [host, distribution, biological control: 314-320]; ZimmerMoHo1986 [biological control: 269-275].

Dactylopius bassi (Targioni Tozzetti)

NOMENCLATURE:

Coccus bassi Targioni Tozzetti, 1866: 129. Nomen nudum; discovered by Ben-Dov & Marotta, 2001: 169.

Coccus bassi Targioni Tozzetti, 1867: 25. Type data: MEXICO: probably on Cactaceae, collected by Dr. Bassi. Syntypes, female. Described: female. Illust. Notes: Type material lost (Giuseppina Pellizzari, Padova, in letter to Yair Ben-Dov).

Dactylopius bassi; Ben-Dov & Marotta, 2001: 169. Change of combination.

DISTRIBUTION: Nearctic: Mexico [Targio1867, BenDovMa2001S].

GENERAL REMARKS: Ben-Dov & Marotta (2001) assigned this species to the Dactylopiidae, with indication that it cannot be distinguished from other species of Dactylopius.

CITATIONS: BenDov2006b [taxonomy, catalogue: 212-213]; BenDovMa2001S [taxonomy, distribution: 169-170]; Targio1866 [taxonomy: 129]; Targio1867 [taxonomy, description, illustration, distribution: 24, 25, 27,]; WilliaBe2009 [taxonomy: 10,12].

Dactylopius ceylonicus (Green)

NOMENCLATURE:

Coccus cacti ceylonicus Green, 1896: 7. Type data: SRI LANKA: Hambantota, on Opuntia sp. Syntypes. Type depository: London: The Natural History Museum, England, UK. Described: female.

Coccus indicus Green, 1908a: 28. Type data: INDIA: Himachal Pradesh, Kangra, on Opuntia dillenii, coll. I. Burkill (Reg. No. 15467). Holotype. Type depository: London: The Natural History Museum, England, UK. Described: female. Synonymy by Green, 1922: 358.

Dactylopius ceylonicus; Sanders, 1909a: 38. Change of combination.

Dactylopius argentinus Dominguez, 1915: 3. Type data: ARGENTINA: States of Cordoba, Santiago del Estero, Rioja and Catamarca, on various species of Cactaceae, such as Opuntia quimilo, O. aurantiaca, O. ficus indica. Syntypes. Described: female. Synonymy by Lizer y Trelles, 1922. Notes: Depository of type material unknown.

Dactylopius indicus; Green, 1922: 358. Change of combination.

COMMON NAME: wild cochineal insect [Green1922].

FOE: COLEOPTERA Coccinellidae: Cryptolaemus montrouzieri Mulsant [TirumaDa1958].

HOSTS: Cactaceae: Cylindropuntia imbricata [ChavezTeCa2011], Nopalea [PerezGKo1992], Nopalea cochenillifera [ChavezTeCa2011], Opuntia [Green1896, DeLott1974a, MacGreSa1984], Opuntia anacantha retrorsa [ClapsDe2001], Opuntia atropes [ChavezTeCa2011], Opuntia canina [DeLott1974a, PerezGKo1992], Opuntia chuquisacana [PerezGKo1992], Opuntia dillenii [Green1908a, Sander1909a, Hutson1926, PerezGKo1992], Opuntia discolor [PerezGKo1992, ClapsDe2001], Opuntia ficus-indica [MacGreSa1984, PerezGKo1992], Opuntia fuliginosa [ChavezTeCa2011], Opuntia hyptiacantha [ChavezTeCa2011], Opuntia longispina [ClapsDe2001], Opuntia monacantha [Green1922, Hutson1926, Ali1967a, DeLott1974a, Varshn1994], Opuntia pilifera [ChavezTeCa2011], Opuntia quimilo [PerezGKo1992, ClapsDe2001], Opuntia retrorsa [DeLott1974a, PerezGKo1992], Opuntia salmiana [ClapsDe2001], Opuntia soehrensii [ClapsDe2001], Opuntia sulphurea [Ferris1919a, PerezGKo1992], Opuntia tomentosa [ChavezTeCa2011].

DISTRIBUTION: Afrotropical: Madagascar [PerezGKo1992]; Mauritania [PerezGKo1992]; Mauritius [Mamet1943a, MoutiaMa1946]; South Africa [DeLott1974a]. Australasian: Australia (New South Wales [PerezGKo1992], Queensland [DeLott1974a, PerezGKo1992]). Nearctic: Mexico (Hidalgo [MacGreSa1984], Jalisco [MacGreSa1984], Mexico State [MacGreSa1984], Morelos [MacGreSa1984], Oaxaca [MacGreSa1984], Veracruz [MacGreSa1984]); United States of America (California [Ferris1919a]). Neotropical: Argentina (Entre Rios [Lizery1922], Formosa [PerezGKo1992], San Luis [Lizery1922, PerezGKo1992], Santa Fe [DeLott1974a, PerezGKo1992], Tucuman [PerezGKo1992]); Bolivia [DeLott1974a, PerezGKo1992]; Brazil [PerezGKo1992]; Paraguay [DeLott1974a, PerezGKo1992]. Oriental: Bangladesh [PerezGKo1992]; India [Green1908a, Sander1909a] (Andhra Pradesh [Varshn1994], Bihar [Ali1967a, Varshn1994], Himachal Pradesh [Varshn1994], Odisha [Varshn1994], Tamil Nadu [Varshn1994], West Bengal [Varshn1994]); Nepal [PerezGKo1992]; Sri Lanka [Green1896, Green1922, PerezGKo1992].

BIOLOGY: Karny (1972) studied in South Africa three populations of Dactylopius spp., namely Species P, Species J and Dactylopius indicus [=Dactylopius ceylonicus]. Studies were carried on life history and their suitability for biological control of Opuntia spp. In Mexico, specimens were collected during April to June on the top portion of their hosts, on ripe cladodes of Opuntia with more than three levels of cladodes and on the areoles of prickly pears, in living fences with scarce vegetation and regosol. In November, D. ceylonicus was collected on the root nodules of Opuntia sp., in wild populations where xerophilous thickets and arenosol predominate. The presence of the insect is scarce without perceptible damage to its host. (Chavez, et al., 2011)

GENERAL REMARKS: Description and illustration of adult female by Ferris (1955a), De Lotto (1974a) and by Perez Guerra & Kosztarab (1992). Description and illustration of nymphs of male and female, and adult male by De Haro (1996). Portillo (1995) distinguished between the fine cochineal species Dactylopius coccus and a group of wild cochineal species comprising D. austrinus, D. ceylonicus, D. confertus, D. confusus, D. opuntiae, D. salmianus, D. tomentosus and D. zimmermani.

STRUCTURE: Colour illustration of the adult female and male given by Green (1922).

SYSTEMATICS: The original description of Dactylopius argentinus Dominguez, 1915 (current status Synonym of D. ceylonicus), includes the binomen, distribution, host plants and the chemical composition of the female. Dactylopius indicus (Green, 1908) has been synonimized with Dactylopius ceylonicus (Green, 1896) by Green (1922). Varshney (2005) presnted an illustration of the adult female of, what he named, Dactylopius indicus Green, 1908, but without further explanations,

ECONOMIC IMPORTANCE AND CONTROL: Goeden (1978) discussed the use of this species in biological control of Opuntia spp.

KEYS: DeHaro & Claps 1995: 16-17 (female) [world]; Perez Guerra & Kosztarab 1992: 35-36 (female) [Species of Dactylopius]; De Lotto 1974a: 173-174 (female) [world]; Ferris 1955a: 86 (female) [North America].

CITATIONS: Ali1967a [host, distribution, life history, economic importance: 30-31]; AndresDaHa1976 [biological control, economic importance: 481]; BadiiFl2001 [host, distribution, economic importance: 503-505]; Beeson1934 [host, distribution, biological control: 203-205]; BenDov2006b [taxonomy, catalogue: 213-214]; Buchne1965 [taxonomy, structure: 296]; Cardon2007 [economic importance, host, distribution: 632-635]; ChavezTeCa2011 [distribution, host: 64]; ChavezTeFr2010 [chemistry, carminic acid: 671-679]; ClapsDe2001 [host, distribution: 79]; DeHaro1996 [taxonomy, description, illustration, host, distribution: 1-4,179-188]; DeHaroCl1995 [taxonomy: 16-17]; DeLott1974a [taxonomy, description, illustration, host, distribution: 175-178]; DennilHo1990 [biological control: 1-10]; DestruPrCo1915 [host, distribution, biological control: 323]; Dodd1927 [host, distribution, economic importance, biological control: 1-44]; Doming1915 [taxonomy, host, distribution, chemistry: 3-5]; Ferris1919a [host, distribution: 20]; Ferris1955a [taxonomy, description, illustration, host, distribution: 90-91]; Frogga1917a [host, distribution, biological control: 417-426]; Goeden1978 [economic importance, biological control, host, distribution: 362-371]; GoedenAn1999 [biological control, host, distribution: 871-890]; Green1896 [taxonomy, description, host, distribution: 7]; Green1908a [taxonomy, description, host, distribution: 28-29]; Green1912a [taxonomy, description, illustration, host, distribution: 80-82,84-87]; Green1922 [taxonomy, description, illustration, host, distribution: 358-359]; Harris2002 [biological control: 74-76]; Hutson1926 [economic importance, biological control, host, distribution: 290-292]; Imms1941 [biological control: 303-305]; Jepson1930 [biological control: 63-72]; Johnst1921 [host, distribution, biological control: 65-66]; Johnst1924 [host, distribution, biological control: 347-401]; Johnst1924a [host, distribution, biological control: 269-290]; Johnst1926 [host, distribution, biological control: 235-240]; Karny1972 [life history, biological control, economic importance, taxonomy, description, illustration, host, distribution: 1-19]; Koszta1987 [economic importance, biological control: 218]; KozarDr1998f [catalogue: 169-170]; LaingHa1976 [biological control: 685]; Lindin1943a [taxonomy: 147]; Lizery1922 [taxonomy, host, distribution: 107-110]; MacGreSa1984 [taxonomy, host, distribution: 217-218]; Mamet1943a [host, distribution: 145]; Mamet1949 [catalogue: 7]; Mann1969 [host, distribution, economic importance, life history: 139-150]; MoranNeHo1986 [biological control: 261-268]; MoranZi1984 [host, distribution, biological control: 297-320]; MoranZi1991 [host, distribution, biological control: 5-27]; MoutiaMa1946 [host, distribution, economic importance: 462]; NanDeWu2013 [phylogenetics: 173]; Ordish1967 [economic importance, biological control]; PerezGKo1992 [taxonomy, description, illustration, host, distribution: 40-44]; Portil1995 [host, distribution: 62-65]; PortilVi2006 [host, distribution, biological control: 249-255]; PulzWoCa2008 [taxonomy: 72]; Ramakr1921a [host, distribution: 343]; Ramakr1931 [economic importance, host, distribution: 229-237]; Reddy1964 [host, distribution, economic importance: 465]; Sander1909a [taxonomy, host, distribution: 38]; Sankar1973 [life history, biological control: 82-88]; SchmutKlLu1957 [host, distribution, economic importance: 426]; Scott1936 [host, distribution, biological control: 404-410]; SerranSoHa2013 [chemistry, economic importance: 6081-6090]; SilvaMeDe2013 [distribution: 5237]; Sulliv1990 [life history, biological control, host, distribution: 123-129]; Sweetm1935 [biological control: 373-377]; Taylor1984 [chemistry, economic importance: 22-25]; TirumaDa1958 [biological control: 545-552]; Tough1938 [host, distribution, biological control: 7-9]; Valles1965 [biological control: 259-279]; Varshn1994 [host, distribution: 331]; Varshn2005 [taxonomy, illustration, host, distribution: 145,147-148]; WilliaBe2009 [taxonomy: 9,12,15,27]; ZimmerMoHo1986 [biological control: 269-275].

Dactylopius coccus Costa

NOMENCLATURE:

Dactylopius cacti auctorum [{non}: Linnaeus, 1758, 1111.

Coccus carti; Modeer, 1778: 44. Misspelling of species name. Notes: Misspelling for Coccus cacti. See Williams & Ben-Dov (2009: 14).

Coccus sativus Lancry, 1791: 486. Type data: SOUTH AMERICA: on Cactaceae. Syntypes, female. Described: female. Synonymy by De Lotto, 1974a: 179. Notes: Regarded as Nomen oblitum by De Lotto (1974a).

Coccus mexicanus Lamarck, 1801: 299. Type data: MEXICO: on Cactaceae. Syntypes, female. Described: female. Synonymy by De Lotto, 1974a: 179. Notes: Regarded as Nomen oblitum by De Lotto (1974a).

Dactylopius coccus Costa, 1829: 16. Type data: ITALY: on Opuntia "Fico d'India". Notes: Type material probably lost (G. Pellizzari in personal information to Yair Ben-Dov). This binomen has been placed on the Official List of Species Names in Zoology. (Name Number 2851) (see ICZN, 1983).

Coccus cacti; Signoret, 1875c: 347. Notes: Signoret named this binomen Coccus cacti des auteurs.

Coccus cacti domestica Meunier, 1884: 19. Synonymy by Lindinger, 1935: 133.

Coccus cacti; Cockerell, 1893dd: 1041.

Pseudococcus signoreti Cockerell, 1900g: 992. Replacement name for Coccus cacti des auteurs, of Signoret, 1875c: 347; synonymy by De Lotto, 1974a: 179.

Dactylopius coccus; Cockerell, 1902k: 454.

Dactylopius mexicanus; Lindinger, 1943c: 249. Change of combination.

Dactylopius sativus; Lindinger, 1943c: 249. Change of combination.

COMMON NAMES: cochineal; cochineal insect; cochineal scale; cochinilla cultivada; cochinilla del carmin; cochinilla fina; grana cochinilla; grana cultivada; grana fina.

FOES: COLEOPTERA Coccinellidae: Chilocorus [PortilVi1998], Exochomus flavipes [RussoMa1996], Hyperaspis fimbriolata Melsheim [Portil2005], Hyperaspis trifurcata Schaeffer [Portil2005], Scymnus hornii LeConte [Portil2005], Scymnus intrusus Horn [Portil2005]. Nitidulidae: Cybocephalus nigritulus LeConte [Portil2005]. DIPTERA Chamaemyiidae: Leucopis bellula Willinston [Portil2005]. LEPIDOPTERA Pyralidae: Laetilia coccidivora (Comstock) [PortilVi1998], Salambona analamprella (Dyar) [Portil2005]. NEUROPTERA Hemerobidae: Sympherobius amiculus Fitch [Portil2005].

HOSTS: Cactaceae: Nopalea cochenillifera [PortilAr1994], Opuntia [DeLott1974a, MacGreSa1984, BenDov2013], Opuntia atropes [PortilAr1994], Opuntia crassa [ChavezTeCa2011], Opuntia ficus-barbarica [PerezGKo1992], Opuntia ficus-indica [MacGreSa1984, PortilAr1994, DeHaroCl1999], Opuntia fuliginosa [ChavezTeCa2011], Opuntia hyptiacantha [MacGreSa1984], Opuntia jaliscana [PortilAr1994], Opuntia megacantha [PortilAr1994], Opuntia pilifera [PortilAr1994], Opuntia sarca [PortilAr1994], Opuntia septacantha [PortilAr1994], Opuntia tomentosa [PerezGKo1992, PortilAr1994], Opuntia vulgaris [Mamet1950].

DISTRIBUTION: Afrotropical: Madagascar [Mamet1943a, Mamet1950]; South Africa [DeLott1974a, PerezGKo1992]. Nearctic: Mexico [PortilAr1994] (Hidalgo [MacGreSa1984], Jalisco [ChavezTeCa2011], Mexico State [ChavezTeCa2011], Oaxaca [MacGreSa1984, PerezGKo1992, Miller1996], Puebla [MacGreSa1984], Veracruz [ChavezTeCa2011]). Neotropical: Argentina (La Rioja [DeHaroCl1999]); Ecuador [PerezGKo1992]; Peru [DeLott1974a, PerezGKo1992, PortilAr1994, Miller1996]; Venezuela [PerezGKo1992]. Palaearctic: Azores [PerezGKo1992]; Canary Islands [Lindin1909c, Lindin1911a, PerezGKo1992, Miller1996, BenDov2013]; Egypt [PerezGKo1992]; Madeira Islands [PerezGKo1992]; Morocco [PerezGKo1992]; Turkey [KaydanUlEr2007].

GENERAL REMARKS: Description and illustration of adult female given by Ferris (1955a), De Lotto (1974a) (while referring to the illustration by Ferris, 1955a) and by Perez Guerra & Kosztarab (1992). Description and illustration of first-instar, second-instar female, second-instar male, prepupa, pupa and adult male by Perez Guerra & Kosztarab (1992). Portillo (1995) distinguished between the fine cochineal species Dactylopius coccus and a group of wild cochineal species comprising D. austrinus, D. ceylonicus, D. confertus, D. confusus, D. opuntiae, D. salmianus, D. tomentosus and D. zimmermani.

STRUCTURE: Its pulverulent white cover and a size larger than the rest of the species of the genus distinguish D. coccus. (Chavez, et al., 2011) Colour photograph of adult female by Foldi (2003b).

SYSTEMATICS: For about 150 years this species was referred to, in scale insect taxonomy, as Coccus cacti Linnaeus, 1758. Cockerell (1899j) demonstrated that the latter species is actually a margarodid. The current status of this species is Protortonia cacti (Linnaeus, 1758) [Margarodidae: Monophlebinae]; see also Ben-Dov (2005). Cockerell selected as a substitute for Coccus cacti the name Dactylopius coccus O. Costa, 1829. Lindinger (1943c) disclosed that Dactylopius coccus O. Costa, was actually a junior synonym of Coccus sativus Lancry, 1791 and Coccus mexicanus Lamarck, 1801. De Lotto (1974a) discussed Lindinger's findings, but remarked that, considering the wide acceptance of Costa's name, Lindinger's (1943c) changes were undesirable. De Lotto (1974a) noted that in order to stabilize Dactylopius coccus, the matter should be referred to the ICZN so that Coccus sativus Lancry, 1791 and Coccus mexicanus Lamarck, 1801, be placed on the Official List of Rejected Names, as nomina oblita. However, no such application has been made. Dactylopius coccus O. Costa, 1829 is accepted in this catalogue, as the currently accepted name of the cochineal insect, while Coccus sativus Lancry, 1791 and Coccus mexicanus Lamarck, 1801, are regarded as nomina oblita that are synonymous with Dactylopius coccus. Dactylopius coccus O. Costa, 1829, has been placed on the Official List of Species Names in Zoology. (Name Number 2851) (see ICZN, 1983).

ECONOMIC IMPORTANCE AND CONTROL: This scale insect was first observed at the XVI century in Mexico by the Spaniards who subsequently introduced it to other Spanish territories as well as to Spain. The body of the adult female of this scale insect is the source of the red carmine dye. It was, and still cultivated for that purpose in Central and South America as well as in the Canary Islands (Donkin, 1977). The insect was, and still is bred for that purpose in Central and South America as well as in the Canary Islands (Donkin, 1977). Portillo & Arreola (1994) recorded a form of Nopalea cochenillifera from Peru resistant to the cochineal. Goeden (1978) discussed the use of this species in biological control of Opuntia spp. Cochineal extract was listed as one of the food ingredients capable of eliciting allergic sensitization, although it would rarely be classified as allergic (Taylor & Hefle, 2001).

KEYS: De Haro & Claps 1995: 16-17 (female) [world]; Perez Guerra & Kosztarab 1992: 35-36 (female) [Species of Dactylopius]; De Lotto 1974a: 173-174 (female) [world]; Ferris 1955a: 86 (female) [North America].

CITATIONS: AbrahaEd1964 [chemistry: 19-25]; AckachPoJa2003 [chemistry: 1028-1034]; AguirrGu1999 [host, distribution, economic importance: 1-389]; AldamaLl2003 [host, distribution, economic importance, life history: 11-19]; AlleviAnBi1998 [host, distribution, chemistry: 575-582]; AlleviAnCi1987 [chemistry: 5469-5472]; AlvarePoVi1996 [life history, economic importance: 19-31]; AquinoBa2002 [structure, chromosomes: 6-10]; AquinoGaCo1994 [structure, chromosomes: 7-23]; Arndt1940 [chemistry, economic importance]; BabithSaPa2003 [chemistry: 258-266]; BalakiVaKa2006 [chemistry: 54-60]; BaranoZySz2004 [chemistry: 54-57]; Barany1978 [economic importance, chemistry, host, distribution, life history: 85-92]; BarberCaIn1972 [host, distribution, economic importance: 10-20]; BarberInPi1995 [host, distribution, economic importance: 1-216]; BarcenAq1997 [life history, host: 21]; BenDov2005a [taxonomy: 263]; BenDov2006b [taxonomy, catalogue: 215-218]; BenDov2013 [distribution, illustration: 72]; Blanch1883 [taxonomy, description, illustration, host, distribution, economic importance, chemistry: 282-302]; Bohmer2000 [chemistry, economic importance: 57-60]; Brana1966 [host, distribution, taxonomy, economic importance: 77-91]; Bustam2002 [economic importance: 21-23]; CalvoSa2000 [chemistry: 439-443]; CamposLl1999 [life history, economic importance]; CamposLl2003 [host, distribution, economic importance, life history: 143-155]; Cardon1999 [economic importance, chemistry: 18-29]; Cardon1999a [economic importance, chemistry: 50-55]; Cardon2007 [taxonomy, economic importance, life history, host, distribution: 607-666]; Castil1993 [host, distribution, life history, ecology, economic importance: 1-94]; ChavezTeCa2011 [distribution, host: 62,64-65]; ChavezTeFr2010 [chemistry, carminic acid: 671-679]; ChenLiBe2004 [life history: 301-303]; ClapsDe2001 [host, distribution: 80]; Cocker1899j [taxonomy: 259-275]; Cocker1900g [taxonomy: 991, 992]; Cocker1902k [taxonomy: 454]; Comsto1881a [taxonomy, description, host, distribution: 346]; ConstaStSa2002 [chemistry: 203-220]; Costa1835 [taxonomy, description, illustration, host, distribution: 16, 17]; CruzDiCrDo1994 [life history, host, distribution: 125-130]; CruzHeGaRo2003 [chemistry: 37-45]; CuevaIzCr2001 [chemistry, economic importance: 144]; Dalrym1984 [chemistry, economic importance: 3-7]; Decary1930 [host, distribution, biological control, ecology: 101-117]; DeHaroCl1995 [taxonomy: 16-17]; DeHaroCl1999 [host, distribution: 128]; DeLott1974a [taxonomy, description, host, distribution: 179, 180]; DeNobr1848 [host, distribution, life history, economic importance: 342-348]; DiazSoSoNu1999 [host, life history, economic importance: 110-111]; Dodd1927 [host, distribution, economic importance, biological control: 1-44]; Donkin1977 [taxonomy, economic importance, life history, host, distribution: 1-84]; DragoMaSa2002 [chemistry: 1969-1974]; Eastwo1984 [chemistry, economic importance: 9-19]; Edelst1963 [chemistry: 15-17]; FAO2001 [chemistry, economic importance: 1-107]; FerreiFeSa2003 [host, distribution, life history: 1560-1568]; FerrerMaAn2005 [chemistry: 243-250]; Ferris1955a [taxonomy, description, illustration, host, distribution: 86-87]; Flores1995 [host, distribution, ecology, life history, economic importance: 35-41]; Flores2002 [host, distribution, life history, ecology: 16-20]; Foldi2003b [taxonomy: 28]; FordGoGr1987 [chemistry: 897-902]; Franci2002 [chemistry: 173-206]; FranciMa2002 [chemistry: 297-330]; FranciWa2002 [chemistry, economic importance: 173-206]; FrancoRuMa2011 [distribution: 7,23]; GarzaZVa1999 [host, life history, economic importance: 112]; Goeden1978 [economic importance, biological control, host, distribution: 362-363]; GoldenBuSt2000 [host, distribution, life history]; GonzalMeCa2002 [chemistry, economic importance: 6968-6974]; Gram1984 [host, distribution, economic importance: 1-299]; GrantGaCa1987 [chemistry: 913-917]; Green1912a [taxonomy, description, illustration, host, distribution: 82-84]; Greenf2004 [history, economic importance: 1-338]; Halpin1996 [chemistry: 76]; Harris2001 [host, distribution, biological control]; Harris2002 [biological control: 74-76]; Haude1998 [chemistry: 240-270]; Hernan1999 [economic importance: 105-118]; HernanDeRo2003 [chemistry: 37-45]; Herrer1983 [host, distribution, economic importance: 26-30]; HeShDe2003 [host, distribution, life history: 604-609]; Hordes1980 [host, distribution, economic importance: 1-237]; ICZN1983 [taxonomy: 77-80]; KarapaCh2006 [chemistry: 75-84]; KarapaDaTs2005 [chemistry: 739-749]; KatoHaTs2004 [chemistry: 241-251]; KaydanUlEr2007 [host, distribution: 93]; KingDu1998 [chemistry: 847-853]; KlugeZiCi1986 [biological control: 295-303]; Koszta1987 [economic importance: 218]; KozarDr1998f [catalogue: 170]; KozielSa2001 [distribution: 1-5]; KreiteMaDi1998a [life history, economic importance: 201-206]; LaGrCo1963 [host, distribution, life history, economic importance: 1-327]; Lamarc1801 [taxonomy, description, host, distribution: 299]; LambdiAqGr2002 [chemistry: 11-15]; Lancry1791 [taxonomy, description, host, distribution: 486]; LeesonAl1999 [chemistry]; Legget1944 [economic importance, chemistry: 1-95]; Lillie1979 [economic importance, chemistry: 57-61]; Lindin1909c [taxonomy, host, distribution: 450]; Lindin1911a [host, distribution: 7]; Lindin1943a [taxonomy: 147]; Lindin1943c [taxonomy: 249-250]; Linnae1758 [taxonomy, description, host, distribution: 457]; LizasoMoGa2000 [chemistry: 549-552]; LlandeCa1999 [host, economic importance: 61-68]; LlandeNi1999 [host, distribution, economic importance]; LlandeNi1999a [host, distribution, life history: 23-29]; MacGreSa1984 [taxonomy, host, distribution: 218-219]; Mamet1943a [host, distribution: 145]; Mamet1950 [host, distribution: 19]; Mann1969 [host, distribution, economic importance, life history: 139-150]; MarinLCi1977 [taxonomy, description, illustration, host, distribution, life history: 115-120]; MarinLCi1983 [host, distribution, life history, ecology: 81-83]; Martog1992 [chemistry: 1-226]; Massch1967 [chemistry: 1080-1085]; MatallCaMi2002 [host, distribution, economic importance, ecology: 22-25]; MazaBaNiLl1999 [life history, chemistry]; McClel2004 [host, distribution: 69-74]; Mendez1999 [host, distribution, life history: 69-78]; MendezGa1999 [host, life history, economic importance: 31-59]; MendezGoLo2004 [chemistry: 1331-1337]; MendezPaTi2003 [economic importance: 165-171]; MendezVeBr1995 [host, distribution, ecology, life history, economic importance: 42-47]; Miller1996 [catalogue, distribution: 69]; MohammGh2008 [distribution: 150]; MoraBl1999 [host, distribution, economic importance: 137]; MoranNeHo1986 [biological control: 261-268]; NanDeWu2013 [phylogenetics: 173]; Nobel1995 [host, distribution, ecology, life history, economic importance: 77-83]; Pelham1963 [host, distribution, taxonomy, economic importance: 635-639]; PerezG1991 [host, distribution, taxonomy, life history: 1-181]; PerezGKo1992 [taxonomy, description, illustration, host, distribution, life history, economic importance, chemistry: 15-49]; PerezS1999 [host, distribution, economic importance: 1-22]; PerezS1999a [economic importance: 119-136]; Pimien1997 [host, distribution, economic importance: 87-95]; PimienNeMu1995 [host, distribution, ecology, life history, economic importance: 1-308]; Portil1992 [host, distribution, economic importance: 50-55]; Portil1993 [life history, economic importance: 25-34]; PortilAr1994 [host, distribution: 90-95]; PortilFl1991 [host, distribution, economic importance: 77-87]; PortilVi1998 [biological control: 43-49]; PortilVi1998a [host, distribution, economic importance: 1-6]; PortilVi1998b [host, distribution, economic importance: 1-21]; PortilVi2002 [host, distribution, economic importance: 249-251]; PortilVi2006 [host, distribution, biological control: 249-255]; PortilViTr1994 [chemistry, economic importance: 38-42]; PowellHo1979 [taxonomy, host, distribution]; PuchalOrAc2003 [chemistry: 1252-1258]; PulzWoCa2008 [taxonomy: 72]; RamireLlRa2008 [morphology, anatomy: 1-5]; ReusHoSt2000 [chemistry: 1836-1839]; Robles1999 [host, life history, ecology: 113-114]; RodrigFaNi2005 [pheromone, life history: 305-306]; RodrigFaSe2005 [chemistry, chemical ecology: 323-329]; RodrigMeNi2001 [life history, ecology: 73-77]; RodrigNi2000 [taxonomy, host, distribution: 85-89]; RodrigPo1989 [host, distribution, economic importance: 49-53]; RubioMe2001 [chemistry]; RubioSaNo2001 [chemistry]; RubioSaNo2001 [chemistry]; RussoMa1996 [life history, distribution, economic importance: 10-13]; RussoMaSu2001 [host, distribution, life history: 333-338]; Saltzm1986 [chemistry, economic importance: 1-84]; Saltzm1992 [chemistry, economic importance: 474-481]; SaltzmKeCh1963 [chemistry, economic importance: 241-251]; Sandbe1994 [host, distribution, economic importance: 1-215]; SarkanMeEv1961 [economic importance, chemistry: 39-40]; SchmutKlLu1957 [host, distribution, economic importance: 426]; Schwep1979 [chemistry: 14-23]; Schwep1986 [chemistry, economic importance: 153-174]; SchwepRo1986 [economic importance, life history, chemistry, host, distribution: 255-283]; SerranSoHa2013 [chemistry, economic importance: 6081-6090]; Signor1869 [taxonomy: 846]; Signor1875c [taxonomy, description, host, distribution: 347-349]; Silva1995 [host, distribution, economic importance: 1-316]; SilvaMeDe2013 [economic importance, distribution: 5237]; SmithMiDa2005 [chemistry: 1010-1030]; Speigh2001 [distribution: 1-5]; Sternl1980 [host, economic importance: 17-21]; SteuriFe2001 [chemistry: 66-72]; Sweetm1935 [biological control: 373-377]; TabarPAlAc2003 [chemistry: 415-419]; Taylor1984 [chemistry, economic importance: 22-25]; Taylor1986 [chemistry: 53-61]; TaylorHe2001 [chemistry, economic importance: 64-69]; Tekele1995 [host, distribution, ecology, life history, economic importance: 48-55]; Tekele1995a [host, distribution, ecology, life history, economic importance: 56-61]; Tekele2001 [host, distribution, economic importance, ecology: 1-191]; Thomso1971 [chemistry: 1-5]; TiedemYa1995 [chemistry: 195-206]; TovarPaGa2005 [host, distribution, economic importance, ecology: 3-7]; TovarPPaMe1999 [host, life history, economic importance: 115-116]; Turok1996 [economic importance: 91-94]; UematsHiSu2002 [chemistry: 335-342]; Valles1965 [biological control: 259-279]; Valles1997 [host, distribution, economic importance: 1-143]; ViguerPo1995 [host, distribution: 24-25]; ViguerPo1995a [chemistry: 41-48]; ViguerPo1997 [life history, economic importance, host, distribution: 135-136]; ViguerPo1999 [host, distribution, life history, ecology: 79-92]; ViguerPo1999a [economic importance: 93-104]; Waller1986 [chemistry, economic importance: 145-155]; Walton1984 [chemistry, economic importance: 30-34]; Warren1914 [host, distribution, description: 387-391]; Willia1969a [taxonomy: 324]; Willia2007b [taxonomy: 441, 427-490]; WilliaBe2009 [taxonomy: 12,14,19,31,42]; Wood1986 [economic importance: 4-8]; Woodwo1903 [taxonomy: 31]; Wouter1985 [chemistry: 119-128]; WouterVe1989a [chemistry: 189-200]; XieXuZh2004a [chemistry: 512-518]; YangYaLi2001 [host, distribution, economic importance: 85-89]; Yannak2003 [distribution, economic importance: 1-5]; ZhangShXu2002 [economic importance: 719-726].

Dactylopius confertus De Lotto

NOMENCLATURE:

Dactylopius confertus De Lotto, 1974a: 180. Type data: ARGENTINA: Salta, Morillo, on Cleistocactus sp. Holotype female. Type depository: Pretoria: South African National Collection of Insects, South Africa. Described: female. Illust.

HOSTS: Cactaceae: Cereus aethiops [ClapsDe2001], Cleistocactus [DeLott1974a], Cleistocactus baumannii [PerezGKo1992, ClapsDe2001], Denmoza rhodocantha [ClapsDe2001], Echinopsis leucantha [PerezGKo1992, ClapsDe2001], Gymnocalycium monvillei [ClapsDe2001], Harrisia tortuosa [ClapsDe2001], Pilocereus [ClapsDe2001], Trichocereus candicans [ClapsDe2001].

DISTRIBUTION: Neotropical: Argentina (Formosa [PerezGKo1992], Salta [DeLott1974a], San Luis [PerezGKo1992]).

GENERAL REMARKS: Description and illustration of adult female by De Lotto (1974a) and by Perez Guerra & Kosztarab (1992). Portillo (1995) distinguished between the fine cochineal species Dactylopius coccus and a group of wild cochineal species comprising D. austrinus, D. ceylonicus, D. confertus, D. confusus, D. opuntiae, D. salmianus, D. tomentosus and D. zimmermani.

KEYS: De Haro & Claps 1995: 16-17 (female) [world]; Perez Guerra & Kosztarab 1992: 35-36 (female) [Species of Dactylopius]; De Lotto 1974a: 173-174 (female) [world].

CITATIONS: BenDov2006b [taxonomy, catalogue: 218-219]; ClapsDe2001 [host, distribution: 80]; DeHaroCl1995 [taxonomy: 16-17]; DeLott1974a [taxonomy, description, illustration, host, distribution: 180-181]; PerezGKo1992 [taxonomy, description, illustration, host, distribution: 49-52]; Portil1995 [host, distribution: 62-65].

Dactylopius confusus (Cockerell)

NOMENCLATURE:

Coccus confusus; Cockerell, 1893dd: 1043. Change of combination.

Acanthococcus confusus Cockerell, 1893gg: 373. Type data: USA: New Mexico, Las Cruces, on a plant of Cactaceae. Syntypes, female. Type depository: Washington: United States National Entomological Collection, U.S. National Museum of Natural History, District of Columbia, USA. Described: female.

Coccus cacti confusus; Cockerell, 1896f: 34. Change of status.

Coccus tomentosus confusus; Cockerell, 1898v: 675. Change of status.

Coccus tomentosus newsteadi Cockerell, 1898v: 675. Synonymy by De Lotto, 1974a: 180.

Pseudococcus tomentosus newsteadi; Cockerell, 1899n: 5. Change of combination.

Pseudococcus confusus; Cockerell, 1899n: 6. Change of combination.

Pseudococcus confusus newsteadi; Cockerell, 1899t: 284. Change of combination.

Dactylopius confusus; Cockerell, 1902k: 454. Change of combination.

Dactylopius confusus newsteadi; Cockerell, 1902k: 454. Change of combination.

Coccus confusus capensis Green, 1912a: 91. Type data: SOUTH AFRICA: Cape Province, on Opuntia monacantha; collected Lounsbury. Syntypes, female. Type depository: London: The Natural History Museum, England, UK. Described: female. Illust. Synonymy by De Lotto, 1974a: 180.

Dactylopius greenii Cockerell, 1929: 329. Replacement name for Coccus confusus capensis Green, 1912a; synonymy by De Lotto, 1974a: 180.

Dactylopius newsteadi; Cockerell, 1929: 329. Change of combination.

Dactylopius confuses; Xie et al., 2004a: 514. Misspelling of species name.

COMMON NAMES: California cochineal scale [Gill1993]; california cochineal scale [Gill1993].

FOES: COLEOPTERA Coccinellidae: Hyperaspis trifurcata Schaeffer [GilreaSm1988]. DIPTERA Chamaemyiidae: Leucopis bellula Williston [GilreaSm1988]. HYMENOPTERA Encyrtidae: Epicerchysius thoreauini Girault [Gordh1979]. LEPIDOPTERA Phycitinae: Laetilia coccidivora (Comstock) [GilreaSm1988].

HOSTS: Cactaceae: Cactus [Riddic1955], Cereus versicolor [PerezGKo1992], Echinocactus [MerrilCh1923], Opuntia [Cocker1896f, Ferris1919a, MerrilCh1923, Ferris1955a, MacGreSa1984, GilreaSm1987], Opuntia aurantiaca [PerezGKo1992], Opuntia dillenii [PerezGKo1992], Opuntia discata [PerezGKo1992], Opuntia elata [PerezGKo1992], Opuntia exaltata [PerezGKo1992], Opuntia ficus-barbarica [PerezGKo1992], Opuntia ficus-indica [ChavezTeCa2011], Opuntia fuliginosa [ChavezTeCa2011], Opuntia humifusa [PerezGKo1992], Opuntia hyptiacantha [ChavezTeCa2011], Opuntia jaliscana [ChavezTeCa2011], Opuntia joconostle [ChavezTeCa2011], Opuntia lindheimeri [PerezGKo1992], Opuntia monacantha [DeLott1974a, PerezGKo1992], Opuntia phaeacantha [ChavezTeCa2011], Opuntia polyacantha [Ferris1919a, PerezGKo1992], Opuntia pumila [PerezGKo1992], Opuntia spinulifera [ChavezTeCa2011], Opuntia streptacantha [PerezGKo1992], Opuntia stricta [DeLott1974a, PerezGKo1992], Opuntia tomentosa [PerezGKo1992], Opuntia wetmorei [Lizery1922], Selenicereus [PerezGKo1992].

DISTRIBUTION: Afrotropical: Mauritius [Miller1996]; South Africa [DeLott1974a, PerezGKo1992, Miller1996]; Zimbabwe [Whelan1965]. Australasian: Australia (Queensland [DeLott1974a], Queensland [PerezGKo1992], Victoria [PerezGKo1992]). Nearctic: Canada (Saskatchewan [MawMo1980, PerezGKo1992, Miller1996]); Mexico [MerrilCh1923, Ferris1955a] (Baja California Norte [MerrilCh1923, Ferris1955a, MacGreSa1984]). Nearctic: Mexico (Chihuahua [MacGreSa1984]). Nearctic: Mexico (Distrito Federal [MacGreSa1984], Durango [Miller1996], Guanajuato [MacGreSa1984], Guerrero [MacGreSa1984, Miller1996], Jalisco [MacGreSa1984, Miller1996], Morelos [MacGreSa1984, Miller1996], Nuevo Leon [Miller1996], Oaxaca [MacGreSa1984], Puebla [MacGreSa1984, PerezGKo1992, Miller1996], Sonora [Miller1996], Tamaulipas [MacGreSa1984], Veracruz [ChavezTeCa2011], Zacatecas [ChavezTeCa2011]); United States of America (Arizona [Ferris1919a, MerrilCh1923, Ferris1955a, Miller1996], California [MerrilCh1923, Ferris1955a, Gill1993, Miller1996], Colorado [MerrilCh1923, Cocker1929, Miller1996], Florida [Miller1996], Idaho [Miller1996], Minnesota [Miller1996], Montana [Ferris1919a], Nebraska [Miller1996], Nevada [Miller1996], New Mexico [Cocker1893dd, MerrilCh1923, Cocker1929, Miller1996], New Mexico [PerezGKo1992], New York [Miller1996], Oklahoma [Miller1996], South Carolina [Miller1996], South Dakota [Miller1996], Texas [Ferris1955a, GilreaSm1987], Utah [Ferris1955a, Miller1996], Washington [Miller1996]). Neotropical: Argentina (Mendoza [Lizery1922]); Bahamas [PerezGKo1992, Miller1996]; Cuba [MerrilCh1923]; Haiti [Miller1996] [PerezG2008] [PerezGKo1992, Miller1996]; Peru [PerezGKo1992, Miller1996]. Palaearctic: Algeria [PerezGKo1992, Miller1996].

BIOLOGY: The bionomics in Texas was studied and described by Gilreath & Smith (1987; 1988). Gill (1993) indicates that it prefers the flat-padded Platyopuntia cacti. Chavez, et al. found that in Mexico, D. confusus grows mainly on the cladodes of tree and shrub cactus forms and on their prickly pear fruits in a predominantly desert habitat with scarce vegetation and arenosol, within the altitude range of 1200 m to 2547 m, which is higher than the previously reported altitude.

GENERAL REMARKS: Description and illustration of adult female by Ferris (1955a), De Lotto (1974a), Perez Guerra & Kosztarab (1992) and by Gill (1993). Portillo (1995) distinguished between the fine cochineal species Dactylopius coccus and a group of wild cochineal species comprising D. austrinus, D. ceylonicus, D. confertus, D. confusus, D. opuntiae, D. salmianus, D. tomentosus and D. zimmermani.

STRUCTURE: Colour photograph given by Gill (1993).

SYSTEMATICS: Insects with the typical morphology of this species were designated by this name (De Lotto 1974, P�rez-Guerra & Kosztarab 1992). Chavez, et al. (2011) designated a biotype 1 with the morphological characteristics diagnosed for D. confusus combined with characteristics corresponding to D. salmianus. Field observations show that these insects promote changes in the color of cladodes and fruits, and when the insects are closely gathered at the trunk-stem and stem-fruit joints, these parts are damaged and may detach from the main plant body. The insects were localized within the altitude range of 1654 m to 2773 m.

ECONOMIC IMPORTANCE AND CONTROL: This species has been introduced into five countries as a biological control agent for suppressing Opuntia weeds; became established in Australia (Dodd, 1940); was established for a time in South Africa (Dodd, 1940; Annecke & Neser, 1977). Gilreath & Smith (1988) proved that natural enemies play a role in regulating population level of this scale on Opuntia cladophylls in Texas, USA. Goeden (1978) discussed the use of this species in biological control of Opuntia spp.

KEYS: De Haro & Claps 1995: 16-17 (female) [world]; Gill 1993: 148 (female) [USA, California]; Perez Guerra & Kosztarab 1992: 35-36 (female) [Species of Dactylopius]; De Lotto 1974a: 173-174 (female) [world]; Ferris 1955a: 86 (female) [North America].

CITATIONS: AndresDa1973 [biological control: 11-28]; AnneckNe1977 [biological control, host, distribution: 303-319]; AquinoGaCo1994 [structure, chromosomes: 7-23]; BenDov2006b [taxonomy, catalogue: 219-221]; ChavezTeCa2011 [distribution, host: 64-68]; ChavezTeFr2010 [chemistry, carminic acid: 671-679]; Cocker1893dd [p. 1043]; Cocker1893gg [taxonomy, description, host, distribution: 373]; Cocker1896f [taxonomy, description, host, distribution: 34]; Cocker1898v [taxonomy, description, host, distribution: 675]; Cocker1899n [taxonomy, host, distribution: 5-6]; Cocker1899t [taxonomy: 284]; Cocker1902k [taxonomy : 454]; Cocker1929 [taxonomy, distribution: 329]; CookGu2004 [taxonomy: 444]; DeHaroCl1995 [taxonomy: 16-17]; DeLott1974a [taxonomy, description, illustration, host, distribution: 180-184]; Dodd1927 [host, distribution, economic importance, biological control: 1-44]; Dodd1940 [biological control: 41-45,54-55,96-105]; Ferris1919a [host, distribution: 20]; Ferris1955a [taxonomy, description, illustration, host, distribution: 88, 89]; Fester1943 [chemistry: 195-196]; Gill1993 [taxonomy, description, illustration, host, distribution: 148, 150]; Gilrea1985 [host, distribution, life history, ecology: 1-202]; GilreaSm1987 [life history, host, distribution, biological control: 768-774]; GilreaSm1988 [life history, economic importance, biological control, host, distribution: 730-738]; Goeden1978 [economic importance, biological control, host, distribution: 362, 365, 368]; GoedenAnFr1974 [biological control, host, distribution: 490-495]; Green1912a [taxonomy, description, illustration, host, distribution: 80,89-93]; GullanCo2001 [taxonomy: 91-99]; GullanCo2007 [taxonomy: 413-425]; GwiazdVaDe2006 [phylogenetics: 16]; Hamlin1924a [biological control: 447-460]; HoskinMcMu1988 [host, distribution, biological control: 115-123]; KozarDr1998f [catalogue: 170-171]; LambdiJo2001 [structure, anatomy: 113-117]; Lindin1943a [taxonomy: 147]; Lizery1922 [taxonomy, host, distribution: 110-111]; MacGreSa1984 [taxonomy, host, distribution: 219-220]; Mann1969 [host, distribution, economic importance, life history: 139-150]; MawMo1980 [host, distribution: 208-211]; MerrilCh1923 [taxonomy, description, host, distribution: 279, 280]; Miller1996 [catalogue, distribution: 71]; MohammGh2008 [distribution: 150]; NanDeWu2013 [phylogenetics: 173]; Pember1964 [host, distribution: 689]; PerezG2008 [distribution: 213]; PerezGKo1992 [taxonomy, description, illustration, host, distribution: 52-58]; Pettey1939 [host, distribution, biological control: 413-415]; Pettey1941 [host, distribution, biological control: 133]; Portil1995 [host, distribution: 62-65]; PortilVi2006 [host, distribution, biological control: 249-255]; Riddic1955 [host : 11]; RossHaOk2012 [phylogeny, taxonomy: 199]; SchmutKlLu1957 [host, distribution, economic importance: 426]; SerranSoHa2013 [chemistry, economic importance: 6081-6090]; Smit1952 [taxonomy: 481-482]; Sweetm1935 [biological control: 373-377]; Taylor1984 [chemistry, economic importance: 22-25]; Whelan1965 [host, distribution, economic importance: 67]; Willia1985a [taxonomy: 219]; WilliaBe2009 [taxonomy: 12,17,24,34,46]; XieXuZh2004a [taxonomy, chemistry: 512-518]; ZhangChSh2004 [life history: 484-489]; ZhangShXu2004 [host, distribution: 321-326].

Dactylopius gracilipilus Van Dam & May

NOMENCLATURE:

Dactylopius gracilipilus Van Dam & May, 2012: 35-38. Type data: USA: Texas, Brewster Co., Big Bend National Park, N29�08.069� W103�02.335�, on Corynopuntia schottii, 8/30/2009, by A.R. Van Dam. Holotype female (examined), by original designation. Type depository: Davis: The Bohart Museum of Entomology, University of California, California, USA. Described: female. Illust.

HOST: Cactaceae: Corynopuntia schottii [VanDamMa2012].

DISTRIBUTION: Nearctic: United States of America (Texas [VanDamMa2012]).

GENERAL REMARKS: Detailed description and illustration in Van Dam & May, 2012.

STRUCTURE: Body oval, length each 1.95�2.84mm, widest point of metathorax each 1.12�2.38mm. Segmentation distinct on meso- and metathorax, and abdomen. Antennae six or seven segmented

SYSTEMATICS: Dactylopius gracilipilus is most similar in appearance to D. tomentosus, but they are easily distinguish by (character-states on D. tomentosus in brackets): (i) presence of abundant simple pores (absent or rare), (ii) clusters of wide-rimmed quinquelocular pores with10 or more pores abundant (large clusters of wide-rimmed pores uncommon), and (iii) truncate setae with parallel sides forming medial longitudinal rows, each seta 1.5�2 times as long as wide (setae in medial longitudinal row each less than 1.5 times as long as wide). Both D. gracilipilus and D. tomentosus infest members of the tribe Cylindropuntieae sensu Wallace and Dickie (2002) (Cactaceae). This might suggest a close affinity between these two species but D. gracilipilus appears to be restricted to Corynopuntia, whereas D. tomentosus are found on Cylindropuntia spp.,(Van Dam & May, 2012)

KEYS: Van Dam & May 2012: 35 (female) [Additional couplet to the key to Dactylopius by Guerra & Koszatarab (1992)].

CITATIONS: VanDamMa2012 [description, distribution, host, illustration, structure, taxonomy: 35-37].

Dactylopius opuntiae (Cockerell)

NOMENCLATURE:

Coccus cacti opuntiae Cockerell, 1896f: 35. Type data: MEXICO: Guanajuato, on a species of Cactaceae. Syntypes, female. Type depository: Washington: United States National Entomological Collection, U.S. National Museum of Natural History, District of Columbia, USA. Described: female. Notes: Cockerell (1896f) credited this species to "Licht. MS", but the author should be named Cockerell.

Coccus tomentosus; Cockerell, 1896f: 35. Misidentification; discovered by De Lotto, 1974a: 184.

Dactylopius opuntiae; Cockerell, 1929: 328. Change of combination.

Dactylopius opuntiae; Mann, 1969: 139,146. Notes: Incorrect citation of "Lichtenstein" as author.

COMMON NAME: opuntia cochineal scale [Gill1993].

FOES: COLEOPTERA Coccinellidae: Coccidophilus citricola [LimaDaFa2011], Exochomus sp. [LimaDaFa2011], Zagreus bimaculosus [LimaDaFa2011]. Nitidulidae: Cybocephalus sp. [LimaDaFa2011]. DIPTERA Chamaemyiidae: Leucopis bellula [SpodekBePr2014]. Syrphidae: Salpingogaster sp. [LimaDaFa2011]. LEPIDOPTERA Pyralidae: Laetilia coccidivora [SpodekBePr2014]. NEUROPTERA Hemerobiidae: Sympherobius barberi [SpodekBePr2014].

HOSTS: Cactaceae: Mammillaria tenuis [PerezGKo1992], Nopalea [PerezGKo1992], Nopelea cochenillifera [ChavezTeCa2011], Nopelea karwinskiana [ChavezTeCa2011], Opuntia [DeLott1974a, AnneckBuCo1976, MacGreSa1984], Opuntia ?lindheimeri [DeLott1974a], Opuntia coccinellifera [PerezGKo1992], Opuntia dillenii [PerezGKo1992], Opuntia engelmannii [ChavezTeCa2011], Opuntia ficus-indica [ChavezTeCa2011, SpodekBePr2014], Opuntia fuliginosa [ChavezTeCa2011], Opuntia fuscicaulis [PerezGKo1992], Opuntia hernandezi [MacGreSa1984], Opuntia hyptiacantha [ChavezTeCa2011], Opuntia inermis [DeLott1974a, PerezGKo1992], Opuntia jaliscana [ChavezTeCa2011], Opuntia leucotricha [ChavezTeCa2011], Opuntia littoralis [PerezGKo1992], Opuntia macdougaliana [ChavezTeCa2011], Opuntia megacantha [Pettey1947, Pettey1950, DeLott1974a], Opuntia megacantha [ChavezTeCa2011], Opuntia occidentalis [PerezGKo1992], Opuntia palmadora [PerezGKo1992], Opuntia robusta [ChavezTeCa2011], Opuntia streptacantha [DeLott1974a], Opuntia stropes [ChavezTeCa2011], Opuntia tardospina [DeLott1974a], Opuntia tomentosa [DeLott1974a, MacGreSa1984], Opuntia tuna [PerezGKo1992], Opuntia undulata [ChavezTeCa2011], Opuntia vulgaris [PerezGKo1992], Platyopuntia [PerezGKo1992].

DISTRIBUTION: Afrotropical: Cape Verde [VanHarCoWi1990]; Kenya [DeLott1974a, PerezGKo1992, Miller1996]; Madagascar [Miller1996]; Mauritius [PerezGKo1992, Miller1996]; Reunion [PerezGKo1992, Miller1996, GermaiMiPa2014]; South Africa [Pettey1947, DeLott1974, AnneckBuCo1976, PerezGKo1992, Miller1996]; Zimbabwe [PerezGKo1992]. Australasian: Australia (New South Wales [DeLott1974a, PerezGKo1992]); Hawaiian Islands (Hawaii [Miller1996]). Nearctic: Mexico (Aguascalientes [MacGreSa1984], Baja California Norte [MacGreSa1984, PerezGKo1992, Miller1996], Chihuahua [ChavezTeCa2011], Distrito Federal [MacGreSa1984, Miller1996], Durango [MacGreSa1984, PerezGKo1992, Miller1996], Guanajuato [Cocker1896f], Guerrero [MacGreSa1984], Hidalgo [MacGreSa1984, PerezGKo1992, Miller1996], Jalisco [MacGreSa1984], Mexico State [MacGreSa1984, Miller1996], Michoacan [MacGreSa1984, PerezGKo1992, Miller1996], Morelos [Miller1996], Nayarit [MacGreSa1984], Nuevo Leon [MacGreSa1984], Oaxaca [MacGreSa1984, PerezGKo1992, Miller1996], Puebla [MacGreSa1984], Queretaro [MacGreSa1984], Tamaulipas [PerezGKo1992, Miller1996], Veracruz [MacGreSa1984], Zacatecas [MacGreSa1984]); United States of America (Arizona [PerezGKo1992, Miller1996], California [PerezGKo1992, Gill1993, Miller1996], Texas [PerezGKo1992, Miller1996]). Neotropical: Brazil (Pernambuco [PerezGKo1992, Miller1996]); Jamaica [PerezGKo1992, Miller1996]; Mexico (Chiapas [MacGreSa1984]). Oriental: India [DeLott1974a, Miller1996] (Odisha [PerezGKo1992], Tamil Nadu [PerezGKo1992]); Pakistan [PerezGKo1992, Miller1996]; Sri Lanka [DeLott1974a, PerezGKo1992, Miller1996]. Palaearctic: France [Miller1996, Foldi2001]; Israel [SpodekBePr2014].

BIOLOGY: Gill (1993) indicated that it prefers the Platyopuntia group of flat-pad cacti in the genus Opuntia.

GENERAL REMARKS: Description and illustration of adult female by De Lotto (1974a), Perez Guerra & Kosztarab (1992) and by Gill (1993). Portillo (1995) distinguished between the fine cochineal species Dactylopius coccus and a group of wild cochineal species comprising D. austrinus, D. ceylonicus, D. confertus, D. confusus, D. opuntiae, D. salmianus, D. tomentosus and D. zimmermani.

STRUCTURE: The ultrastructure of wax-secreting glands described and SEM photographed by Hartley et al. (1983). Colour photograph given by Gill (1993).

SYSTEMATICS: Examination of the morphological characteristics of the insects of the species D. opuntiae by Chavez, et al (2011) resulted in two separate groups, designated D. opuntiae and D. opuntiae biotype 1, described as follows. Insects with a typical morphology of this species (De Lotto 1974, P�rez-Guerra & Kosztarab 1992) were designated D. opuntiae. Insects with morphological characteristics of D. opuntiae combined with characteristics corresponding to D. salmianus were designated as Dactylopius opuntiae biotype 1. D. salmianus has a thinner and elongated body and the structures of the setae and pores are more elongated and more separated or dispersed over the insect body. (Chavez, et al, 2011)

ECONOMIC IMPORTANCE AND CONTROL: Pettey (1947), Pettey (1950), Pettey & Marais (1950) and Zimmermann & Moran (1991) presented detailed accounts on the role of this species in the successful biological control of prickly pears, especially Opuntia megacantha, in South Africa. Goeden (1978) discussed the use of this species in biological control of Opuntia spp.

KEYS: De Haro & Claps 1995: 16-17 (female) [world]; Gill 1993: 148 (female) [USA, California]; Perez Guerra & Kosztarab 1992: 35-36 (female) [Species of Dactylopius]; De Lotto 1974a: 173-174 (female) [world].

CITATIONS: Andres1981 [biological control, economic importance, host, distribution: 341-349]; AndresGo1971 [biological control: 143-164]; AnneckBuCo1976 [biological control, economic importance, host, distribution: 111-116]; BalciuBuPu1995 [distribution, biological control: 445]; BenDov2006b [taxonomy, catalogue: 221-222]; Bennet1971 [host, distribution, biological control: 371]; ChavezTeCa2011 [distribution, host: 68]; ChavezTeFr2010 [chemistry, carminic acid: 671-679]; Cocker1896f [taxonomy, distribution: 35]; Cocker1929 [taxonomy: 329]; Coyle1990 [host, distribution, biological control: 490-492]; daSilvDuDa2010 [host, distribution, life history, physiology, ecology: 718-722]; DeHaroCl1995 [taxonomy: 16-17]; DeLott1974 [taxonomy, description, illustration, host, distribution: 184-186]; duToit1942 [host, distribution, economic importance, biological control: 300-304]; FloresMuRu2006 [host, distribution, life history: 97-102]; Foldi2001 [distribution: 303-308]; FoxcroHo2000 [host, distribution, economic importance, biological control, life history: 1-5]; GermaiMiPa2014 [distribution: 22]; Gill1993 [taxonomy, description, illustration, host, distribution: 149, 151]; GithurZiHo1999 [host, distribution, economic importance, biological control, life history, taxonomy: 43-48]; Goeden1978 [economic importance, biological control, host, distribution: 362-378]; GoedenAnFr1974 [biological control, host, distribution: 490-495]; HartleWaMo1983 [structure: 97-98]; Heap1990 [biological control]; HoffmaImVo2002 [host, distribution, economic importance, biological control: 900-908]; HoffmaMoZi1999 [host, distribution, economic importance, biological control: 15-20]; HuffakCa1986 [biological control, economic importance: 95-107]; HuffakDaJa1984 [distribution, biological control: 659-691]; KlugeZiCi1986 [biological control: 295-303]; Koszta1987 [economic importance, biological control: 218]; LaingHa1976 [biological control: 685]; LimaDaFa2011 [biological control: 51-54]; MacGreSa1984 [taxonomy, host, distribution: 220-221]; Mann1969 [host, distribution, economic importance, life history: 139-150]; Miller1996 [catalogue, distribution: 71]; MoranZi1984 [host, distribution, biological control: 297-320]; NanDeWu2013 [phylogenetics: 173]; Ordish1967 [economic importance, biological control]; PalaciNiLl2004 [chemical control: 99-106]; PerezGKo1992 [taxonomy, description, illustration, host, distribution: 59-62]; Pettey1943 [host, distribution, biological control: 743-746]; Pettey1946a [host, distribution, biological control: 31-33]; Pettey1947 [behaviour, ecology, economic importance, biological control, life history, host, distribution: 102-136]; Pettey1950 [life history, ecology, biological control, host distribution: 1-12]; PetteyMa1950 [economic importance, chemical control, host, distribution: 13-34]; Portil1995 [host, distribution: 62-65]; PulzWoCa2008 [taxonomy: 72]; SerranSoHa2013 [chemistry, economic importance: 6081-6090]; SilvaMeDe2013 [molecular data: 5236-5246]; Skaife1979 [host, distribution, life history: 1-3]; SkaifeLeBa1981 [taxonomy, distribution: 1-5]; SpodekBePr2014 [distribution, host: 377�379]; Sweetm1935 [biological control: 373-377]; Valles1965 [biological control: 259-279]; VanHarCoWi1990 [host, distribution: 132]; VolchaHoZi1999 [host, distribution, economic importance, biological control, life history, taxonomy: 85-91]; Weber1951 [host, distribution, biological control: 327]; WilliaBe2009 [taxonomy: 12,35]; Wilson1964 [biological control: 238]; Zimmer1989a [biological control: 1-2]; ZimmerMo1982 [host, distribution, biological control: 314-320]; ZimmerMo1991 [host, distribution, economic importance, biological control: 29-35]; ZimmerMoHo1986 [biological control: 269-275].

Dactylopius salmianus De Lotto

NOMENCLATURE:

Dactylopius salmianus De Lotto, 1974a: 186. Type data: ARGENTINA: Chaco, Santa Sylvina, on Opuntia salmiana. Holotype female. Type depository: Pretoria: South African National Collection of Insects, South Africa. Described: female. Illust.

HOST: Cactaceae: Opuntia salmiana [DeLott1974a].

DISTRIBUTION: Neotropical: Argentina (Catamarca [PerezGKo1992], Chaco [DeLott1974a]).

GENERAL REMARKS: Description and illustration of adult female by De Lotto (1974a) and by Perez Guerra & Kosztarab (1992). Portillo (1995) distinguished between the fine cochineal species Dactylopius coccus and a group of wild cochineal species comprising D. austrinus, D. ceylonicus, D. confertus, D. confusus, D. opuntiae, D. salmianus, D. tomentosus and D. zimmermani.

KEYS: Perez Guerra & Kosztarab 1992: 35-36 (female) [Species of Dactylopius]; De Lotto 1974a: 173-174 (female) [world].

CITATIONS: BenDov2006b [taxonomy, catalogue: 222-223]; ClapsDe2001 [host, distribution: 80]; DeLott1974a [taxonomy, description, illustration, host, distribution: 186-188]; PerezGKo1992 [taxonomy, description, illustration, host, distribution: 65-68]; Portil1995 [host, distribution: 62-65].

Dactylopius tomentosus (Lamarck)

NOMENCLATURE:

Coccus silvestris Lancry, 1791: 486. Type data: SOUTH AMERICA: on Cactaceae. Syntypes, female. Described: female. Synonymy by De Lotto, 1974a: 188. Notes: Regarded a nomen oblitum by De Lotto (1974a).

Coccus tomentosus Lamarck, 1801: 299. Type data: FRANCE: Paris, in greenhouse of the Museum of Natural History, on Cactaceae. Syntypes, female. Described: female. Notes: Depository of type material unknown.

Coccus cacti silvestris Meunier, 1884: 19. Synonymy by Williams & Ben-Dov, 2009: 13.

Acanthococcus tomentosus; Cockerell, 1893dd: 1042. Change of combination.

Pseudococcus tomentosus; Cockerell, 1899n: 5. Change of combination.

Dactylopius tomentosus; Cockerell, 1902k: 454. Change of combination.

Dactylopius silvestris; Lindinger, 1943c: 250. Change of combination.

COMMON NAMES: tomentose cochineal scale [Gill1993].

FOE: COLEOPTERA Coccinellidae: Cryptolaemus montrouzieri Mulsant [PuttarCh1957, TirumaDa1958, BaskarLaUt1999, BaskarSrMu2002].

HOSTS: Cactaceae: Cereus versicolor [MacGreSa1984], Cylindropuntia fulgida fulgida [MathenHoHo2009], Nopalea karwinskiana [ChavezTeCa2011], Opuntia [Ferris1919a, MacGreSa1984], Opuntia acanthocarpa [PerezGKo1992], Opuntia coccinellifera [MacGreSa1984], Opuntia dillenii [Hutson1926], Opuntia ficus-indica [MacGreSa1984], Opuntia fulgida [MacGreSa1984], Opuntia hernandezi [MacGreSa1984], Opuntia imbricata [PerezGKo1992], Opuntia indica [MacGreSa1984], Opuntia leptocaulis [PerezGKo1992], Opuntia megacantha [MacGreSa1984], Opuntia monacantha [Hutson1926], Opuntia tomentosa [MacGreSa1984], Opuntia vulgaris [MacGreSa1984].

DISTRIBUTION: Afrotropical: Madagascar [Frappa1932]; Mauritius [Mamet1943a]; Reunion [Mamet1952, GermaiMiPa2014]; South Africa [Miller1996]. Australasian: Australia (New South Wales [PerezGKo1992, Miller1996]). Nearctic: Mexico (Baja California Norte [MacGreSa1984, Miller1996], Chihuahua [ChavezTeCa2011]). Nearctic: Mexico (Chihuahua [MacGreSa1984]). Nearctic: Mexico (Coahuila [MacGreSa1984], Distrito Federal [MacGreSa1984], Guanajuato [MacGreSa1984], Jalisco [ChavezTeCa2011], Nuevo Leon [MacGreSa1984], Oaxaca [Miller1996]); United States of America (Arizona [Ferris1919a, Miller1996], California [Ferris1919a, Gill1993, Miller1996], Nevada [Miller1996], New Mexico [Miller1996], Texas [Miller1996], Utah [Miller1996]). Neotropical: Argentina (Buenos Aires [Lizery1922, Miller1996]); Colombia [Kondo2001].

BIOLOGY: The insects develop exclusively on the cladodes of their hosts and their tiny size makes them almost imperceptible. They do not damage or promote changes in the plant and develop in a desert habitat where xerophilous thickets predominate, on vertisol and arenosol. The presence of spiders was frequently observed with this species. (Chavez, et al., 2011)

GENERAL REMARKS: Description and illustration of adult female by Ferris (1955a), De Lotto (1974a), Perez Guerra & Kosztarab (1992) and by Gill (1993). Portillo (1995) distinguished between the fine cochineal species Dactylopius coccus and a group of wild cochineal species comprising D. austrinus, D. ceylonicus, D. confertus, D. confusus, D. opuntiae, D. salmianus, D. tomentosus and D. zimmermani.

STRUCTURE: Colour photograph given by Gill (1993).

SYSTEMATICS: Lindinger (1943c) disclosed that Coccus silvestris Lancry, 1791 predated Coccus tomentosus Lamarck, 1801. De Lotto (1974a) discussed Lindinger's findings, but remarked that Lindinger's (1943c) changes were undesirable, and regarded Coccus silvestris Lancry, 1791 a nomen oblitum.

ECONOMIC IMPORTANCE AND CONTROL: Goeden (1978) discussed the use of this species in biological control of Opuntia spp. Mathenge et al. (2009) studied and evaluated the use of this species for biological control of Cylindropuntia fulga in South Africa. Mathenge et al. (2009, 2009a, 2009b, 2010) discussed various apsects of the application of this scale insects in biological control of cactus weeds.

KEYS: Van Dam & May 2012: 35 (female) [Additional couplet to the key to Dactylopius by Guerra & Koszatarab (1992)]; De Haro & Claps 1995: 16-17 (female) [world]; Gill 1993: 148 (female) [USA, California]; Perez Guerra & Kosztarab 1992: 35-35 (female) [Species of Dactylopius]; De Lotto 1974a: 173-174 (female) [world]; Ferris 1955a: 86 (female) [North America].

CITATIONS: Alexan1931 [host, distribution, biological control: 226]; BaskarLaUt1999 [biological control: 605-606]; BaskarSrMu2002 [biological control: 54-56]; BenDov2006b [taxonomy, catalogue: 223-224]; ChavezTeCa2011 [distribution, host: 68-69]; ChavezTeFr2010 [chemistry, carminic acid: 671-679]; Cocker1893dd [taxonomy, description: 1041-1049]; Cocker1898v [taxonomy, description, host, distribution: 675]; Cocker1899n [taxonomy: 5]; Cocker1902k [taxonomy: 454]; Dass1934 [host, distribution, biological control: 342-345]; DeHaroCl1995 [taxonomy: 16-17]; Dodd1927 [host, distribution, economic importance, biological control: 1-44]; Drea1973 [biological control: 49-55]; Ferris1919a [host, distribution: 20]; Ferris1955a [taxonomy, description, illustration, host, distribution: 92-93]; Frappa1932 [host, distribution, biological control: 48-55]; GermaiMiPa2014 [distribution: 22]; Gill1993 [taxonomy, description, illustration, host, distribution: 149, 152]; Goeden1978 [economic importance, biological control, host, distribution: 362]; Graham1960 [host, distribution, biological control: 40]; Green1912a [taxonomy, description, illustration, host, distribution: 79,87-89]; Hamlin1924 [host, distribution, life history, biological control: 60-64]; Hamlin1926 [host, distribution, life history, biological control: 97-105]; Hamlin1932 [host, distribution, biological control, economic importance: 89-120]; HenderZi2003 [host, distribution, biological control: 46-47]; Hutson1926 [economic importance, biological control, host, distribution: 290-292]; Imms1931 [biological control: 124-127]; Jepson1930 [biological control : 63-72]; Kondo2001 [taxonomy, host, distribution: 40]; KozarDr1998f [catalogue: 171]; Lamarc1801 [taxonomy, description, host, distribution: 299]; Lancry1791 [taxonomy, description, host, distribution: 486]; LeelaDMu1953 [host, distribution, economic importance, biological control: 219-224]; Lindin1943a [taxonomy: 147]; Lindin1943c [taxonomy: 250]; Lizery1922 [taxonomy, host, distribution: 110]; MacGreSa1984 [taxonomy, host, distribution: 222-223]; Mamet1943a [host, distribution: 145]; Mamet1949 [catalogue: 7-8]; MathenHoHo2009 [economic importance, biological control, life history, host, distribution: 551-559]; MathenHoHo2009a [economic importance, biological control, life history, host, distribution: 619-627]; MathenHoHo2009b [economic importance, biological control, life history, host, distribution: 1-12]; MathenHoHo2010 [economic importance, biological control, life history, host, distribution: 1-8]; Miller1996 [catalogue, distribution: 71]; MoranZi1991 [host, distribution, biological control: 37-55]; MoutiaMa1946 [host, distribution, economic importance: 462]; NanDeWu2013 [phylogenetics: 173]; Painte1951 [economic importance, control]; PerezGKo1992 [taxonomy, description, illustration, host, distribution: 68-72]; Portil1995 [host, distribution: 62-65]; PortilVi2006 [host, distribution, biological control: 249-255]; PuttarCh1957 [biological control: 7]; Ramach1931 [economic importance, host, distribution, biological control: 14-15]; Reddy1964 [host, distribution, economic importance: 465]; SchmutKlLu1957 [host, distribution, economic importance: 426]; Signor1869 [taxonomy: 873]; Taylor1984 [chemistry, economic importance: 22-25]; TirumaDa1958 [biological control: 545-552]; Tooke1930 [host, distribution, biological control: 386-393]; UsmanPu1955 [host, distribution: 47]; VanDamMa2012 [taxonomy: 35]; WilliaBe2009 [taxonomy: 13,43,44,46]; ZimmerMoHo1986 [biological control: 269-275].

Dactylopius zimmermanni De Lotto

NOMENCLATURE:

Dactylopius zimmermanni De Lotto, 1974a: 189. Type data: ARGENTINA: Mendoza, Vargas, on Tephrocactus ovatus. Holotype female. Type depository: Pretoria: South African National Collection of Insects, South Africa. Described: female. Illust.

HOSTS: Cactaceae: Cereus aethiops [ClapsDe2001], Maihueniopsis darwinii [ClapsDe2001], Maihuentia patagonica [ClapsDe2001], Tephrocactus ovatus [DeLott1974a].

DISTRIBUTION: Neotropical: Argentina (Mendoza [DeLott1974a, ClapsDe2001]).

GENERAL REMARKS: Description and illustration of adult female by De Lotto (1974a) and by Perez Guerra & Kosztarab (1992). Portillo (1995) distinguished between the fine cochineal species Dactylopius coccus and a group of wild cochineal species comprising D. austrinus, D. ceylonicus, D. confertus, D. confusus, D. opuntiae, D. salmianus, D. tomentosus and D. zimmermani.

KEYS: De Haro & Claps 1995: 16-17 (female) [world]; Perez Guerra & Kosztarab 1992: 35-36 (female) [Species of Dactylopius]; De Lotto 1974a: 173-174 (female) [world].

CITATIONS: BenDov2006b [taxonomy, catalogue: 224]; ClapsDe2001 [host, distribution: 80]; DeHaroCl1995 [taxonomy: 16-17]; DeLott1974a [taxonomy, description, illustration, host, distribution: 189-191]; PerezGKo1992 [taxonomy, description, illustration, host, distribution: 72-76]; Portil1995 [host, distribution: 62-65].

Species Removed from Family

Amonostherium hymenocleae