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Coccidology. the Study of Scale Insects (Hemiptera: Sternorrhyncha: Coccoidea)

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Coccidology. the Study of Scale Insects (Hemiptera: Sternorrhyncha: Coccoidea) View metadata, citation and similar papers at core.ac.uk brought to you by CORE 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 insects (Hemiptera: Takumasa Kondo1, Penny J. Gullan2, Douglas J. Williams3 Sternorrhyncha: 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 years 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 insect 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, coccidae, 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 taxonomy is based mainly de insectos escama. C entomology 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 Carl Linnaeus’ 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 species 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); “Coccus”, the genus 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 thelytoky and haplodiploidy (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. Margarodidae “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 Diaspididae changes that have happened in the and Eriococcidae). 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 ants 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 ant-scale association in Macaranga plants has been a subject 1 Entomologist Ph.D. Corporación Colombiana de Scale insects are sap 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 aphids (Aphidoidea), whiteflies have symbiotic relationships with stingless CA 95616, U.S.A. 3 Entomologist Ph.D. Department of Entomology, The (Aleyrodoidea) and jumping plant lice bees (Hymenoptera: Apidae: Meliponinae) Natural History Museum, London SW7 5BD, U.K. (Psylloidea), 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 insect 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 family to (1923) reported that the Mexicans used the known only as fossils. Scale insects are which they belong, e.g., the armoured lac of Tachardiella fulgens Comstock, under generally divided into two informal groups, scales (Diaspididae), the mealybugs the name of “jomilla”, medicinally and the archaeococcoids and the neococcoids. (Pseudococcidae), the putoids (Putoidae), 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 wax from Ceroplastes africanus Green which are absent in the neococcoids. The (Margarodidae), lac insects (Kerriidae), in Egypt is used for welding porcelain archaeococcoids consist of 27 families, cochineal insects (Dactylopiidae), and and mending metal cracks and holes. i.e., 15 extant families (Callipappidae, ensign scales (Ortheziidae). The most Carayonemidae, Coelostomidiidae, commonly encountered families are Another scale insect, Kermes vermilio Kuwaniidae, Marchalinidae, Margarodidae, those with the most species, namely Planchon (Kermesidae), that produces a Matsucoccidae, Monophlebidae, the Diaspididae, Pseudococcidae and red dye, has been known for more than Ortheziidae, Phenacoleachiidae, Coccidae. two millennia and lives on species of Pityococcidae, Putoidae, Steingeliidae, oaks around the Mediterranean shores Stigmacoccidae and Xylococcidae) and 12 Although among the ortheziids (Foldi, 2003). These insects were originally fossil 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 Newsteadia 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 phloem-sap feeders; however, some feed Porphyrophora hameli Brandt that lives The neococcoids are composed of on parenchyma tissue by directly feeding mainly on grass roots in Armenia and 17 extant families, i.e., Aclerdidae, on the contents of parenchymatic cells. surrounding countries was widely used Asterolecaniidae, Beesoniidae, Scales are found on various parts of for dying silks (Donkin, 1977b). A related Cerococcidae, Coccidae, Conchaspididae, their hosts, and may infest leaves, twigs, insect Porphyrophora polonica (L.) found in Dactylopiidae, Diaspididae, Eriococcidae, branches and roots, and some live inside Poland and surrounding areas, known as Halimococcidae, Kermesidae, Kerriidae, plant domatia. Some scale insects are even the Polish cochineal insect, also feeding Lecanodiaspididae, Micrococcidae, known to survive on plants completely on roots, was widely used to produce a Phoenicococcidae, Pseudococcidae and submerged at high tide (Harrison, 1916). red dye and exported to Western Europe. Stictococcidae; 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 instars 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 toxins, transmitting viruses John Philip Breyn (Breyn, 1731) showed however, we consider this family (and the or excreting honeydew, which serves as how an insect could be
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