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provided by Ciencia y Tecnología Agropecuaria (E-Journal) Revista Corpoica – Ciencia y Tecnología Agropecuaria (2008) 9(2), 55-61

Review article Coccidology. The study of scale (: Takumasa Kondo1, Penny J. Gullan2, Douglas J. Williams3 : Coccoidea)

Coccidología. El estudio de insectos ABSTRACT escama (Hemiptera: Sternorrhyncha: A brief introduction to the science of coccidology, and a synopsis of the history, Coccoidea) advances and challenges in this field of study are discussed. The changes in coccidology since the publication of the Systema Naturae by Carolus Linnaeus 250 ago are RESUMEN Se presenta una breve introducción a la briefly reviewed. The economic importance, the phylogenetic relationships and the ciencia de la coccidología y se discute una application of DNA barcoding to scale identification are also considered in the sinopsis de la historia, avances y desafíos de discussion section. este campo de estudio. Se hace una breve revisión de los cambios de la coccidología Keywords: Scale, insects, , DNA, history. desde la publicación de Systema Naturae por Carolus Linnaeus hace 250 años. También se discuten la importancia económica, las INTRODUCTION Sternorrhyncha (Gullan & Martin, 2003). relaciones filogenéticas y la aplicación de These insects are usually less than 5 mm códigos de barras del ADN en la identificación occidology is the branch of in length. Their is based mainly de insectos escama. C that deals with the study of on the microscopic cuticular features of hemipterous insects of the superfamily Palabras clave: insectos, escama, coccidae, the adult female. The adult female is ADN, historia. Coccoidea, particularly on areas related paedomorphic, maturing in a juvenile to systematics. For the purpose of this form, whereas the adult male (when synopsis, we set the starting point for the present), after going through a prepupal study of coccidology as 1758, beginning and pupal stage, turns into an alate with with ’ 10th edition of the non-functional mouthparts. The Coccoidea Systema Naturae (Linnaeus, 1758). During form a rather small group of insects this period of 250 years, the number of in terms of richness with some described scale insects has increased from 7,700 species described. However, scale 24 species (Williams, 2007) to some 7,700 insects are an interesting group of insects species in more than 1,050 genera (Ben- to study. According to Gullan & Cook Dov et al., 2006). The root of the word (2007), scale insects have great variation coccidology is derived from the word in chromosome number (Nur et al., 1987); “”, the in which Linnaeus sperm structure (Robison, 1977; 1990); included the bulk of his scale insects. Most types of bacterial endosymbioses (Buchner, scale insects were not recognisable as 1965; Thao et al., 2002; Gruwell et al., 2005, insects by the ancients, but rather as seeds 2007); and genetic systems, including or berries, and were given the ancient hermaphroditism, diplodiploidy, Greek word “Kokkos” and then the later and (Nur, 1980; Latin word “Coccus” meaning a berry. Normark, 2003). Their morphology The word “coccidology”, as a branch of varies greatly amongst members of the entomology, was probably coined for the different families, with some species first time by Tinsley (1899) in his article producing cysts (e.g. “Contributions to coccidology. I.” Here sensu stricto) that can live underground we attempt to summarize briefly how for many years, and other species are coccidology has changed in the last 250 highly modified to live under the bark years, with emphasis on the remarkable of their hosts (e.g., some changes that have happened in the and ). For ecologists and field in the XXI century. This account evolutionary biologists, scale insects are supplements a brief history of Coccoidea often subjects to study because of their by Ferris (1957). mutualistic relationships with tending Radicado: 15 de septiembre de 2008 and their close associations with their hosts. Aceptado: 9 de diciembre de 2008 What are scale insects? For example, the -scale association in Macaranga plants has been a subject 1 Entomologist Ph.D. Corporación Colombiana de Scale insects are sucking hemipterous of studies in Southeast Asia (Heckroth Investigación Agropecuaria Corpoica, C.I. Palmira, Colombia. [email protected] insects that include all members of the et al. 1998; Ueda et al., 2008). Moreover, 2 Entomologist Ph.D. Department of Entomology, superfamily Coccoidea. These are closely some scale insects are even known to University of California, One Shields Avenue, Davis, related to (Aphidoidea), have symbiotic relationships with stingless CA 95616, U.S.A. 3 Entomologist Ph.D. Department of Entomology, The (Aleyrodoidea) and jumping plant lice (: : Meliponinae) Museum, London SW7 5BD, U.K. (), which make up the suborder (Camargo & Pedro, 2002).

© 2008 Corporación Colombiana de Investigación Agropecuaria 56 Coccidology. The study of scale insects (Hemiptera: Sternorrhyncha: Coccoidea)

There are currently 46 known Scale insects are known by various centuries. In the New World, Chamberlin families, of which 32 are extant and 14 are names depending on the to (1923) reported that the Mexicans used the known only as . Scale insects are which they belong, e.g., the armoured of Tachardiella fulgens Comstock, under generally divided into two informal groups, scales (Diaspididae), the the name of “jomilla”, medicinally and the archaeococcoids and the neococcoids. (Pseudococcidae), the putoids (), for repairing crockery and other utensils. The archaeococcoids are defined by the the soft scales (Coccidae), the felt Kamel & Afifi (1970) have reported how presence of 2–8 pairs of abdominal spiracles, scales (Eriococcidae), ground pearls the from africanus Green which are absent in the neococcoids. The (Margarodidae), lac insects (), in Egypt is used for welding porcelain archaeococcoids consist of 27 families, insects (Dactylopiidae), and and mending metal cracks and holes. i.e., 15 extant families (Callipappidae, ensign scales (). The most Carayonemidae, Coelostomidiidae, commonly encountered families are Another scale insect, vermilio Kuwaniidae, Marchalinidae, Margarodidae, those with the most species, namely Planchon (), that produces a , , the Diaspididae, Pseudococcidae and red , has been known for more than Ortheziidae, , Coccidae. two millennia and lives on species of Pityococcidae, Putoidae, Steingeliidae, around the Mediterranean shores Stigmacoccidae and ) and 12 Although among the ortheziids (Foldi, 2003). These insects were originally families (Electrococcidae, Jersicoccidae, Arctorthezia cataphracta (Olafsen) is known thought to be little worms, hence the Latin Kukaspididae, Labiococcidae, Naibiidae to feed on a basidiomycete fungal species name vermiculi from which the name and seven recently described families, (Thorpe, 1968) and kanayana vermilion is derived (and similar names namely Arnoldidae, Lithuanicoccidae, Kawai & Takagi on fungal mats (Kawai, in languages derived from Latin). Weitschatidae, Grohnidae, Serafinidae 1980), the majority of scale insects feed (Koteja, 2008), and Hammanococcidae and on plants, especially flowering plants Armenian red, a name for the red Lebanococcidae (Koteja & Azar, 2008)). (angiosperms). Scale insects are generally dye obtained from the scale insect -sap feeders; however, some feed hameli Brandt that lives The neococcoids are composed of on parenchyma tissue by directly feeding mainly on grass roots in and 17 extant families, i.e., , on the contents of parenchymatic cells. surrounding countries was widely used , , Scales are found on various parts of for dying (Donkin, 1977b). A related , Coccidae, , their hosts, and may infest , twigs, insect Porphyrophora polonica (L.) found in Dactylopiidae, Diaspididae, Eriococcidae, branches and roots, and some live inside and surrounding areas, known as , Kermesidae, Kerriidae, plant domatia. Some scale insects are even the insect, also feeding , Micrococcidae, known to survive on plants completely on roots, was widely used to produce a , Pseudococcidae and submerged at high tide (Harrison, 1916). red dye and exported to Western Europe. ; and 2 extinct families, Many are important pests of agriculture Both of these insects are peculiar in that namely Inkaidae and the recently (e.g., Peronti et al., 2001; Miller et al., the intermediate encyst and these described Pennygullaniidae (Koteja 2005; Culik et al., 2007) and may injure cysts were originally thought to be of plant & Azar, 2008). Koteja and Azar (2008) or kill plants by depleting them of their origin. A famous treatise on this insect by considered the Putoidae a neococcoid, sap, injecting , transmitting viruses John Philip Breyn (Breyn, 1731) showed however, we consider this family (and the or excreting , which serves as how an insect could be studied in detail probably related Labiococcidae) to belong a medium for sooty moulds (Williams and illustrated showing all the instars. to the archaecoccoids. The adult females of & Granara de Willink, 1992; Gullan & species of Putoidae superficially are most Martin, 2003). In the New World, the cochineal similar to those of the neococcoid family insect of commerce, coccus Pseudococcidae, with most females of Scale insects have been known for Costa, a species used by Mayans, Aztecs these families possessing trilocular pores, centuries, not just for the damage they and Incas, interested many European cerarii and dorsal ostioles. However, the cause, but for the useful red that some workers. The red dye produced by this adult females of Putoidae differ from of them produce, for valuable secretions insect proved to be superior to any of the those of Pseudococcidae in having: (i) in the form of and resins, and even red dyes produced by other scale insects three or four campaniform sensilla on for their use as medicine and food. In the (for an account see Donkin, 1977a) and each surface of each trochanter; (ii) three Oriental Region, Mahdihassan (1954), for at a time the species was even grown pairs of interflagellar setae; and (iii) a pair instance, has given us an account of how on in North Africa. The Spanish of basal denticles on each claw (Hardy et lac insects were known to the Chinese in also exported supplies to southern Asia al., 2008). Furthermore, the adult males of writings dating to 320 AD when the insects via the Philippines from South America putoids differ from those of pseudococcids produced a red dye and a substance for (Donkin, 1977b). Currently, the cochineal in many morphological features (Hardy et sticking things together. Lac insects are insect is grown for commercial purposes al., 2008; Hodgson & Foldi, 2006) and now known to be tropicopolitan. The most in Chile, Mexico, Peru and the Canary the chromosome system of putoids does important species are still deliberately Islands. Cochineal dye is still produced not show paternal genome elimination, grown in India and surrounding countries commercially in chemical factories in which is characteristic of neococcoids for and sealing wax. The use of Europe and the USA (Pérez Guerra & (Cook et al., 2002). lac in India probably dates back many Kosztarab, 1992) and has been used as

Revista Corpoica – Ciencia y Tecnología Agropecuaria (2008) 9(2), 55-61 Coccidology. The study of scale insects (Hemiptera: Sternorrhyncha: Coccoidea) 57 a biological control agent of noxious 2007). The most recent redescription of C. only a few of his mica slides have survived weeds (Moran & Zimmermann, hesperidum by Hodgson (1994) consists of (Deyrolle, 1875). 1984; Volchansky et al., 1999). three and a half pages beginning with a section on classification and nomenclature, There then followed important works Scale insects provide other products a description including the morphology based on slide-mounted specimens by too. It is generally thought that flower of the insect in life and of slide-mounted William Miles Maskell in New Zealand nectar is the main ingredient of . specimens as seen under a compound from 1879–1897 and by Theodore Dru However, honeybees collect other sweet microscope, a figure, a section on the Alison Cockerell, Edward Ernest Green, ingredients, especially when flowers are material studied and a discussion section Robert Newstead and many French scarce. In Greece and Turkey, honeybees where the author considers its affinities workers whose slide collections are collect honeydew from Marchalina with other coccids. In Linnaeus’ time, a housed in the United States National hellenica (Gennadius) (Marchalinidae) single sentence sufficed to describe C. Museum of Natural History (actually feeding on trees and, in Greece hesperidum, however, the family Coccidae at the USDA, Beltsville, Maryland), alone, this “” accounts in which C. hesperidum is included The Natural History Museum, London, for 60%–65% of all honey produced currently contains more than 1,100 species and the Muséum National d’Histoire (Hodgson & Gounari, 2006). In Middle in more than 100 genera! Even after 250 Naturelle, Paris. Although identifications Europe, about 50% of all honey produced years, C. hesperidum continues to be a are possible from many illustrations by is from honeydew, particularly from common scale insect in greenhouses, but these authors, we owe much gratitude the soft scale hemicryphus we know now that there are many other to Gordon Floyd Ferris who, in a series (Dalman), a species found mostly on scale insects that are commonly found in of articles published in the journal the Norway spruce, Picea abies (Kunkel, greenhouses. For example, in southeastern Microentomology from 1936–1955 and 1997). According to Kunkel (1997), the USA, according to Baker (1994), the list of in his Atlas of Scale Insects of North honeydew of many species of scale common scale insects in greenhouses and America (1937–1955), adopted a method insects in at least six families are known on indoor plants includes the mealybugs: of illustration first used by Karl Šulc as a source of honey worldwide. citri (Risso), (Šulc, 1895). This method includes a full longispinus (Targioni Tozzetti), outline of the insect divided by a line in The pela wax scale, Ericerus pela gossypii Townsend & Cockerell [probably a the middle and showing the dorsum on (Chavannes), is cultivated in China misidentification of Ph. madeirensis Green], the left and the venter on the right with for the production of high quality wax falcifer Kunckel d’Herculais, the enlargements of important characters (Qin, 1997). The wax of this soft scale is armoured scales boisduvalii Signoret either elsewhere on the illustration produced by the immature stages (second- and aspidistrae (Signoret), and or around the perimeter of the main nymphs, prepupa and pupae) of the soft scales, L., drawing. This unique drawing method for the males and is used for making candles, Eucalymnatus tessellatus (Signoret) and scale insects was adopted later by Alfred coating material for pills, papers, and coffeae (Walker). Thus, Linnaeus’s Serge Balachowsky and most subsequent for shining leather products and tires 1758 description of C. hesperidum is not authors and has stood the test of time amongst its uses. useful in the present day. so that accurate identifications can now be made from printed works. Although Some scale insects are also used as It was Réné Antoine Ferchault de access to slide collections is necessary, it human food. In Australia, aboriginal Réaumur who produced a remarkable has even become clear that some of the people eat the -inducing scale, account of scale insects, mainly of oldest microscope slide preparations in Cystococcus pomiformis (Froggatt), which Europe, with illustrations of the external collections can be remounted successfully according to the natives has a watery appearance (Réaumur, 1738), many of when necessary. female and nutty-flavoured nymphs which can be recognised today. Linnaeus (Gullan & Cranston, 2005). In Sakorn (1758), in his Systema Naturae, the starting Scale insects as economic pests Nakorn Province, Thailand, the giant point of zoological nomenclature, drew , Nietnera sp. (Margarodidae heavily on Réaumur’s work for his There have probably been outbreaks of sensu lato) is cooked together with sticky chapter on the genus Coccus. scale insects causing damage to local rice and consumed (Kondo, 2001). crops and plants for centuries, but the In the 18th and 19th centuries, some of arrival in the USA of purchasi History of Coccidology the European countries were interested in Maskell (Monophlebidae) towards the the fauna of their overseas territories. Insects end of the 19th century, resulting in the Like many other fields in entomology, were sent to Europe for identification and almost collapse of the industry, coccidology has gone through its own it became clear that identification from the seemed to attract attention throughout the evolution. The original Latin description of external appearance was not satisfactory. world. Outbreaks causing considerable Coccus hesperidum (L.), as given in Systema With better microscopes, entomologists damage are occurring to the present day Naturae (1758: 455-457) can be fitted in one such as Signoret in many works on scale and in the last 40 years the accidental sentence as follows: “The Coccus of the insects from 1860–1886 produced articles introduction of the mealybug greenhouses; It lives on evergreen trees” based on slide-mounted specimens (Ben- Matile-Ferrero (English translation as given in Williams, Dov & Matile-Ferrero, 1995). Unfortunately, (Pseudococcidae) from South America to

Revista Corpoica – Ciencia y Tecnología Agropecuaria (2008) 9(2), 55-61 58 Coccidology. The study of scale insects (Hemiptera: Sternorrhyncha: Coccoidea)

West Africa caused considerable damage link a DNA sequence to a certain species, archaeococcoid families occurred well to cassava throughout Africa affecting the which needs to be identified by the prior to that of extant families. staple food of 200 million people (Herren traditional taxonomic expert. They suggest the lack of resolution of & Neuenschwander, 1991). About the relationships among scale insect families same time, another mealybug Rastrococcus The classification of the Coccoidea, from 18S data and morphology indicates invadens Williams was introduced to West particularly the archaeococcids has gone that the basal radiations might have been Africa from the Oriental Region affecting through a series of overhauls in the last 40 relatively rapid (Gullan & Cook, 2007). a wide variety of fruit trees (Agounké years. Koteja (1974) introduced a multi- et al., 1988). In more recent times, the family classification for scale insects In to elucidate the unresolved mealybug hirsutus based on the morphology of mouthparts phylogenetic relationships of scale insects, (Green) was introduced accidentally to the recognizing a number of families formerly future studies may consider increasing Caribbean area affecting a large number included in the Margarodidae sensu lato. taxon sampling and the number of infor- of plant species including fruit trees and In a special edition of the online journal mative genetic markers, as well as adding plants of economic importance (Chang Zootaxa celebrating the 300th anniversary more morphological data, especially from & Miller, 1996). Yet another mealybug of the birth of Carolus Linnaeus (Zhang & the adult males and first-instar nymphs. species, the mealybug Shear, 2007), Gullan & Cook (2007) gave marginatus Williams & Granara de Willink a summary of our current understanding Insects of the suborder Sternorrhyncha that had been known only locally in of the higher classification of the harbor maternally transmitted bacteria Mexico, suddenly spread to much of the Coccoidea. Many coccidologists now housed in a specialized organ called Caribbean area and beyond (Miller et accept that the superfamily Coccoidea the bacteriome (von Dohlen et al., al., 2001). All of these species were comprises up to 32 extant families. The 2001). Mealybugs have primary and brought under control by or neococcoids are considered more derived secondary endosymbionts belonging predators with the aid of taxonomists who than the archaeococcoids and form a to different subdivisions of the phylum could identify the pest species accurately monophyletic group supported by both Proteobacteria, although some mealybug and suggest areas where natural enemies morphological and genetic data, but the species lack secondary endosymbionts could be located. More recently, the lobate monophyly of the archaeococcoids is (von Dohlen et al., 2001; Thao et al., 2002). lac scale Paratachardina pseudolobata Kondo uncertain and their higher-level ranks The primary endosymbionts of armoured & Gullan has caused serious damage in have been controversial. scale insects belong to a different phylum, Florida, the Bahamas and in Christmas the Bacteroidetes, and their phylogeny Island, Australia (Kondo & Gullan 2007; Recent studies using the nuclear small follows closely that of their scale insect Schroer et al., 2008). In Florida alone, the subunit ribosomal RNA gene (SSU rRNA hosts (Gruwell et al., 2007). This appears lobate lac scale has been recorded on over or 18S) have helped resolve some of the also to be the case in mealybugs in which 300 species of plants (Howard et al., 2006). higher relationships within the Coccoidea, the mealybug microbial ecology appears Now this scale has also been reported in particularly those of the neococcoids. strongly correlated with phylogeny Cuba, and no effective natural enemies According to Gullan and Cook (2007), future (Downie & Gullan, 2005; Hardy et al., have been found because its place of studies may show that some of the species- 2008). Perhaps it may be possible to help origin is still unknown. poor families are autapomorphic members elucidate unresolved relationships of scale of a larger group, e.g., the Aclerdidae and insects by looking at the relationships of One would expect that these outbreaks Micrococcidae are similar to Coccidae and their endosymbionts. might lead to an increase in workers their recognition at family rank may render studying the group but in three of the the Coccidae paraphyletic; the Beesoniidae, DNA Barcoding and scale insects world’s most important centres for scale Dactylopiidae and Stictococcidae are each insects housed in the USDA at Beltsville, closely related to a different subset of the DNA barcoding uses a short fragment Maryland, The Natural History Museum, Eriococcidae in molecular phylogenetic of the mitochondrial DNA to link an London, and the Natural History Museum studies; the Phoenicococcidae is monotypic organism to a species (Herbert & Gregory, in Paris, there are no full-time employed and together with the Halimococcidae 2005). The mitochondrial DNA (mtDNA) incumbents and research is carried out by has affinities to the Diaspididae; the of eukaryote cells has a relatively fast retired associates or collaborators. Conchaspididae resemble the diaspidids mutation rate, resulting in significant and also other distantly related variance in mtDNA sequences between The phylogenetic relationships of scale neococcoids, and its phylogenetic position species but generally small variation within insects is an enigma. species. The concept of barcoding has been successfully implemented in organisms New tools and resources are making the The higher-level relationships of the such as some birds (Hebert et al., 2004a), study of scale insects by taxonomists more archaeococcoids still remain unresolved. mammals (Borisenko et al., 2008), and exciting, and in some administration centres According to Gullan and Cook (2007), insects, especially (e.g., Hebert it is thought that eventually identifications the extent of 18S divergence among the et al., 2004b). The genetic marker of choice from the DNA of a species could dispense coccoid families is as high as or higher for DNA barcoding (sanctioned by the with the skill of the traditional taxonomist. than among the aphid families, and Consortium for the Barcode of Life, CBOL) However, there will always be a need to it seems that the radiation of extant has been the 5’ region of the CO1 gene,

Revista Corpoica – Ciencia y Tecnología Agropecuaria (2008) 9(2), 55-61 Coccidology. The study of scale insects (Hemiptera: Sternorrhyncha: Coccoidea) 59 but in scale insects, universal primers fail al., 2006), which allows users to obtain Borisenko AV, Lim BK, Ivanova NV, Hanner to amplify the region for any but a few information on numerous aspects of most RH, Hebert PDN. 2008. DNA barcoding in taxa. Instead the D2 region of 28s rDNA described species by just the click of a surveys of small mammal communities: a field study in Suriname. Molecular Ecology is currently being proposed by scale insect button. Advances in molecular genetics Resources 8(3): 471-479. workers as a possible marker for barcoding are helping to resolve the phylogenetic of scale insects. The barcoding method in relationships of this morphologically Breyn JP. 1731. Historia naturalis Cocci Radicum Tinctorii quod plonicum vulgo coccidology is closely associated with the highly derived group, and new techniques audit. Gedani, viii, 22 p. preparation of voucher specimens and the such as barcoding are being contemplated correct identification of the scale insect by as a tool for identification of common Buchner P. 1965. Endosymbiosis of with Plant Microorganisms. New York, taxonomic experts. The aim of barcoding in pest species. Despite all of these advances Interscience Publishers / John Wiley, 909 p. scale insects is the accurate, economic and in technology, and our accumulated fast identification of important pest species. knowledge about scale insects, the field Camargo JMF, Pedro SRM. 2002. Mutualistic Association between a tiny Amazonian The damage caused by scale insects is said of coccidology still faces many challenges. The higher-level relationships of scale stingless bee and a wax-producing scale to sum up to billions of dollars in damage insect. Biotropica 34(3): 446-451. and control every (Kosztarab, 1990). insects are far from being resolved, and Chamberlin JC. 1923. A systematic monograph Coccidologists in general agree that DNA every year there are new species of of the Tachardiinae or lac insects (Coccidae). barcoding is not a replacement tool for scale insects being added to the list of agricultural pests. Perhaps, the greatest Bulletin of Entomological Research 14:147- systematics, but rather a tool that can be 212. used for identification of morphologically problem that the field of coccidology faces today is the decline of scale insect Chang LWH, Miller CE. 1996. Pathway Risk defined species. We agree with Kipling specialists worldwide. Museums that used Assessment. Pink mealybug from the and Rubinoff (2004) in that an extremely to employ coccidology experts no longer Caribbean. In: USDA-APHIS, Planning and well developed background knowledge risk analysis systems, policy and program replace the retirees. We only hope that of the taxa to be sampled and an a priori development, Washington DC, 61 p. the science of coccidology will continue understanding of sequence variation to progress. Cook LG, Gullan PJ, Trueman HE. 2002. A among populations and individuals preliminary phylogeny of the scale insects are needed in order to use properly the (Hemiptera: Sternorrhyncha: Coccoidea) barcoding method. ACKNOWLEDGEMENTS based on nuclear small-subunit ribosomal DNA. Molecular Phylogenetics and Evolution 25(1): 43-52. Many thanks to Dr. Lyn G. Cook, University of Queensland, for reviewing Culik MP, Martins DS, Ventura JA, Peronti the barcoding section of the manuscript. ALBG, Gullan PJ, Kondo T. 2007. Coccidae, Pseudococcidae, Ortheziidae, and Monophlebidae (Hemiptera: Coccoidea) of Thanks also to Juan Jaramillo Vásquez, Espírito Santo, Brazil. Biota Neotropica 7(3): Corpoica, Centro de Investigación 61-65. Palmira, and three anonymous reviewers Deyrolle AME. 1875. Les collections for kindly reviewing the manuscript. entomologiques. La collection d’Héniptères de M. Signoret. Petites Nouvelles Entomologiques 121:483-484. REFERENCES Donkin RA. 1977a. Spanish red - an Figure 1. DNA Barcoding and scale insects Agounké D, Agricola U, Bokonon-Ganta ethnographical study of cochineal and HA. 1988. Rastrococcus invadens Williams the Opuntia cactus. Transactions of the (Hemiptera: Pseudococcidae), a serious American Philosophical Society 67(5): 1-84. DISCUSSION exotic pest of fruit trees and other plants in West Africa. Bulletin of Entomological Donkin RA. 1977b. The insect dyes of western and west-. Anthropos 72: 847- The advances in coccidology during the Research 78:695-702. 880. last 250 years have been remarkable. Baker JR. 1994. 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