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Ciencia y Tecnología Agropecuaria

Print version ISSN 0122-8706

Corpoica cienc. tecnol. agropecu. vol.14 no.1 Mosquera Jan./June 2013





A new species of Toumeyella Cockerell (Hemiptera: Coccoidea: Coccidae) on coffee roots, Coffea arabica L. (Rubiaceae), from Colombia and Venezuela


Una nueva especie de Toumeyella Cockerell (Hemiptera: Coccoidea: Coccidae) en raíces de café, Coffea arabica L. (Rubiaceae) en Colombia y Venezuela


Takumasa Kondo1

1 Entomologist PhD. Corporación Colombiana de Investigación Agropecuaria (Corpoica), Centro de Investigación Palmira, Calle 23, Carrera 37, Palmira, Valle, Colombia,

Fecha de recepción: 12-03-2013. Fecha de aceptación: 11-04-2013


A new species of soft scale insect from Colombia and Venezuela, Toumeyella coffeae Kondo sp. nov., is described and illustrated based on the adult female. A taxonomic key to the genera of scale insects of the Toumeyella-group and a list of 49 soft scale insect (Coccidae) species recorded from Coffea spp. worldwide are provided.

Key words: coccid, scale insect, soft scales, Toumeyella coffeae.


Una nueva especie de escama blanda de Colombia y Venezuela, Toumeyella coffeae Kondo sp. nov., se describe e ilustra con base en la hembra adulta. Se proveen una clave taxonómica para separar los géneros de escamas blandas del grupo-Toumeyella y una lista de 49 especies de escamas blandas (Coccidae) registradas sobre Coffea spp., en el mundo.

Palabras claves: cóccido, insecto escama, escama blanda, Toumeyella coffeae.



Williams and Kondo (2008) gave an overview of 16 species in the genus Toumeyella Cockerell, providing information about the distribution, host plants and comments on the morphology of some species. Later, Kondo & Williams (2009) erected the genus Neotoumeyella into which they transferred Toumeyella cerifera Ferris and T. sonorensis Ckll. & Parrott as Neotoumeyella cerifera (Ferris) and N. sonorensis (Ckll. & Parrott), and more recently Kondo & Pellizzari (2011) described Toumeyella fontanai from Mexico. Currently, there are 15 named species of soft scale insects included in the genus Toumeyella, distributed in Brazil (2 spp.), Cuba (1 sp.), Mexico (5 spp.) and the United States (9 spp.) (Kondo & Pellizzari, 2011). There are further undescribed species recorded from Cuba on Adelia ricinella L. (Euphorbiaceae), Casearia sylvestris Sw. (Salicaceae), Coffea arabica (Rubiaceae) and Eugenia foetida Pers. (Myrtaceae) (Mestre et al., 2011) and from Colombia on avocado, Persea americana L., on Booth 7, Booth 8, Choquette, Lorena, Santana and Trinidad cultivars (Kondo et al., 2011). There is another undescribed species of Toumeyella, known in the literature as Toumeyella sp. that has been considered for a long time as a pest of coffee in Venezuela, where it affects the roots and root crown of its host (Barrera, 2008; Fernández, 1957; Le Pelley, 1968; Murphy, 1997). The species described here from specimens from Colombia and Venezuela, collected also from coffee roots may correspond to the species reported above as a pest of coffee in Venezuela, however, this needs confirmation. On the other hand, the species reported on coffee in Cuba comes from the aerial parts of the plant and the morphology is different from the new species herein described (N. Mestre-Novoa, personal communication).

The main purpose of this paper is the description of a new species of Toumeyella collected from the roots of Coffea arabica plants. The adult female of this new species is described and illustrated. A taxonomic key to separate the genera included in the Toumeyella-group (excluding Neolecanium that is considered a synonym of Toumeyella (Kondo, 2004b, 2009) is provided for easy identification of closely related genera that occur in Central and South America. Furthermore, a list of soft scale insects (Hemiptera: Coccoidea: Coccidae) reported on Coffea spp. (Rubiaceae) worldwide is provided as an aid to quarantine personnel and researchers working on scale insect pests.



Specimens were slide-mounted following the procedure described by Williams & Granara de Willink (1992). The terminology of morphological features follows mostly that of Kondo & Williams (2008) and Kondo & Pellizzari (2011). Measurements are given as a range and were taken from 10 specimens (the best two slidemounted specimens for each of the five collecting sites, including the holotype). Measurements of the holotype are given in parenthesis. The range of body size was determined by selecting the smallest and the largest specimens in the material studied. The illustration of the adult female (Fig. 2) shows the dorsum on the left side and the venter on the right side with enlargements of important features around the margin. The material studied is deposited in the following institutions:

UNAB: Museo Entomológico Facultad de Agronomía, Universidad Nacional de Colombia, sede Bogotá, Bogotá, Cundinamarca, Colombia.
UVCO: Museo de Entomología, Universidad del Valle, Cali, Valle del Cauca, Colombia.

A list of soft scale insects (Hemiptera: Coccoidea: Coccidae) reported on Coffea spp. worldwide was compiled mostly from information found in the scale insect database ScaleNet (Ben-Dov et al., 2013) with additional information from the literature. The valid name of each species was checked in ScaleNet.




Key to genera of the Toumeyella-group based on adult females (adapted from Kondo, 2010a; Kondo & Williams, 2009):

1. Anal plates located near mid-dorsum; anterior spiracular pore band incomplete, not extending to margin (posterior spiracular pore band extending to margin); marginal setae cylindrical, peg-like .............. ............................................. Cyclolecanium Morrison

- Anal plates not located near mid-dorsum, usually found at about 1/5 to 1/4 of body length from posterior margin; anterior and posterior spiracular pore bands complete, both pore bands extending to margin; marginal setae variable, not peg-like .......................... 2

2. Dorsal microducts around body margin conspicuously larger than rest of microducts on dorsum .................................................. Octolecanium Kondo

- Dorsal microducts around body margin not conspicuously larger than rest of microducts on dorsum ................................................................................. 3

3. Preopercular pores absent; dorsum with dense pattern of invaginated bilocular microducts; stigmatic spines absent ................................ Pseudophilippia Cockerell

- Preopercular pores present; dorsal microducts variable, generally without dense pattern of invaginated bilocular microducts; stigmatic spines usually present ............................................................................................ 4

4. Ventral tubular ducts absent.................................... 5

- Ventral tubular ducts present at least around perivulvar area ................................................................................ 6

5. Dorsum of slide-mounted specimens with a dense pattern of microducts. Young adult females in life covered by a white cottony to powdery wax, although this powdery wax disappears in old specimens. Perivulvar pores mostly with 5 loculi and one central loculus ................................................... Bombacoccus Kondo

- Dorsum of slide-mounted specimens without a dense pattern of microducts. Adult females in life covered by a thin layer of glassy wax. Perivulvar pores mostly with 7 or 8 loculi and with 2 or 3 central loculi ......... ......................................................... Akermes Cockerell

6. Ventral tubular ducts present at least in a submarginal band on abdominal region and reaching area around posterior spiracular pore band ................................. ............................. Neotoumeyella Kondo and Williams

- Ventral tubular ducts not distributed as above ............ ........................................................................................ 7

7. Ventral tubular ducts located around vulva and extending anteriorly in a medio-lateral line up to about abdominal segment V; dorsal setae lanceolate; stigmatic clefts deep.................... Megasaissetia Cockerell

- Ventral tubular ducts located around vulva and often also on posterior abdominal segments, but not extending anteriorly in a medio-lateral line; dorsal setae generally sharply spinose, rarely lanceolate; stigmatic clefts shallow or absent ............................ .................................................. Toumeyella Cockerell

Toumeyella coffeae Kondo, sp. nov. (Figures 1 & 2)

Proposed common name: English: Coffee-root scale. Spanish: Escama blanda de la raíz del café.

Material studied. Holotype. Adult female (UNAB). Colombia: Norte de Santander: municipio de Cucutilla, Vereda Pedregal Bajo, Finca Buenos Aires, 2.v.2013, coll. Manuel Alexis Leal, ex roots of Coffea arabica. Paratypes. Same data as holotype 31 slides (28 adult females + 3 third-instar nymphs) (UNAB). Municipio Arboledas, vereda Aguadas, Finca El Porvenir, 2.v.2013, coll. Fabio Alonso Jáuregui, ex roots of Coffea arabica, 36 (28 adult females + 28 first-instar nymphs + 8 male second-instar nymphs + 4 female second-instar nymphs + 3 female third-instar nymphs) (UNAB). Municipio Cucutilla, Finca La Dalia, 07,34513 °N, 072,47227 °W, 1,497 m a.s.l., 10.iv.2013, coll. María Fernanda Díaz Niño, ex café Canastilla, 20 (26 adult females + 8 first-instar nymphs) (UNAB). Municipio Arboledas, vereda La Rinconada, Finca El Cedro, 1,526 m a.s.l., 25.xi.2012, coll. Clemencia Villegas G., ex roots of Coffea arabica var. Colombia, 20 (15 adult females + 17 first-instar nymphs + 1 female third-instar nymph) (UVCO).

Other material studied. Venezuela: Estado de Lara: Sanare, Sabana Redonda Arriba, 1,100 m a.s.l., vi.2010, coll. Radamés Urtiaga, #1255, 2 (2 adult females + 1 first-instar nymph + 1 female second-instar nymph + 1 female third-instar nymph) (UNAB).

Description. Adult female (Figures 1 & 2)

Unmounted material. Body convex. Derm orange, brown to greyish green in color, with dark tesselations, but usually with mid-dorsum very lightly or not mottled; anal plates dark to reddish brown. Mature insects 1.5- 4.2 mm in diameter (Figure 1).

Mounted material. Body outline oval to elongate oval, often narrowing anteriorly; body 1.7-5.3 (1.8) mm long, 1.1-4.1 (1.4) mm wide (Figure 2).

Dorsum. Derm membranous on young adult females, becoming heavily sclerotized in older specimens. Dorsal setae (dset) sharply spinose, straight or slightly curved, each 8.6-20.0 (9-17) mm long, rather scarse, more or less scattered evenly. Dorsal microducts (dmic) each 2.0-2.5 mm wide, with a long terminal filament, evenly scattered. Simple pores (sp) each about 2.2-3.0 mm wide, evenly scattered. Dorsal tubular ducts absent. Preopercular pores (prop) present on mid-dorsum anterior to anal plates, extending anteriorly up to area just dorsad to labium, each pore 5-14 (6-11) mm wide. Dorsal tubercles and pocket-like sclerotizations absent. Anal plates (aplt) together quadrate, plates located at about 1/5 to 1/4 of body length from posterior margin, each plate 167-203 (183-191) mm long, 93- 118 (98-110) mm wide, anterolateral margin 123-164 (125-142) mm long, posterolateral margin 113-147 (140-142) mm long, with 1 or 2 setae on dorsal surface, plus 1 pair of long fringe setae, about 7 ventral subapical setae and 5 pairs of hypopygial setae. Anal ring with 10 setae (setae represented by dark dots). A small sclerotic area often present just anterior to anal plates.

Margin. Marginal setae (mset) sharply spinose, straight to slightly bent, each 12-50 (17-30) mm long, arranged in a single, often irregular row, with 5-10 (6 on holotype) on each side between anterior and posterior stigmatic areas. Stigmatic clefts well developed, usually with 3 spines per stigmatic area; stigmatic spines (stgsp) bluntly to sharply spinose, median stigmatic spine usually longest, 29-44 (27-42) mm long, lateral spines 15-22 (13-36) mm long. Eyes not detected.

Venter. Derm entirely membranous. Ventral setae slender, straight or slightly bent, each 6-27 mm long; also 3 pairs of long median setae, each 23-70 mm long, a pair on last segment (VII) longest. Ventral microducts (vmic) scattered evenly throughout, each about 3 mm wide. Ventral tubular ducts scarce, present around vulvar region, each tubular duct with a terminal filament ending in a small, branched terminal gland, duct opening 4-5 mm wide. Perivulvar pores (pvp) rather scarce, small, about same size as spiracular pores, each 5-6 mm wide, with 4-6 loculi (mostly with 5 loculi), rarely a few up to 8 loculi or more, present around vulvar area, a few on abdominal segment VII, and in small groups of about 1-6 pores submedially on each abdominal segment and lateral to metathoracic leg. Spiracular pores (spp) each 5-6 mm wide, with 4-6 loculi, mostly with 5 loculi, rarely a few pores with more loculi, present in a narrow band 1-3 pores wide, rarely up to 6 in some sections, extending laterally from each spiracle to body margin. Spiracles well developed, anterior spiracular peritremes each 56- 78 (61-65) mm wide, posterior peritremes each 66-88 (70-71) mm wide. Legs greatly reduced, segments not discernible, fused, except for claw, total length of all legs including claw, each 59-105 (80-102) mm long, metathoracic legs usually largest; claws well developed, without a denticle, claw digitules, slender, knobbed; tarsal digitules knobbed, longer than claw digitules.

Antennae (ant) short, each 44-100 (73-84) mm long, segmentation poorly developed, 2-4 (2) segmented, with fleshy setae present on apex of antennae. With 3 pairs of interantennal setae, each 5-32 (5-20) mm long. Mouthparts well developed, clypeolabral shield 167- 240 (240) mm wide; labium 1 segmented, with 4 pairs of labial setae.

Etymology. The species is named after its Type host: coffee.

Notes. First-instar nymphs of T. coffeae have fivesegmented antennae and resemble other species of Toumeyella (Kondo & Williams, 2002; Sheffer & Williams, 1996; Williams, 1993).

Biology. This species is sexual because second-instar males have been found in one infestation. These coccids live and feed on roots and on underground parts of stems. In the collection from Finca Buenos Aires at Vereda Pedregal Bajo, Norte de Santander, Colombia, on 2.v 2013, Toumeyella coffeae was found together with the mealybug Dysmicoccus brevipes (Cockerell) (Pseudococcidae).

Soft scale insects (Coccidae) on coffee, Coffea spp. in the world

According to the scale insect database ScaleNet (Ben- Dov et al., 2013) there are 47 species of soft scale insects recorded on Coffea spp. worldwide, excluding species identified only to genus level and Lecanium mercarae Ramakrishna Ayyar, which is a nomen nudum (Ben- Dov, 1993). A total of 49 species of soft scale insects on Coffea spp., including the new species, T. coffeae, are listed here. Table 1 gives the country or countries from which each species has been recorded on Coffea spp., additional country records not listed in ScaleNet (Ben-Dov et al., 2013) and the validation source. Toumeyella liriodendri (Gmelin) listed by Barrera (2008) on the roots of coffee in Guatemala was omitted from Table 1 because this is likely a misidentification since T. liriodendri is a North American species that feeds on the aerial parts of its host (Kondo & Williams, 2008; Gill, 1988; Hamon & Williams, 1984). Coccus asiaticus Lindinger recorded by Le Pelley (1968) and cited by Murphy (1997) is a synonym of Parasaissetia nigra (Nietner), thus its distribution is listed under P. nigra



The soft scales of the genus Toumeyella belong to the subfamily Myzolecaniinae (Hodgson 1994) as well as the Toumeyella-group (Steinweden, 1929). The Myzolecaniinae was erected by Hodgson (1994) to include species that lack a number of structures of the cuticle and have reduced appendages. The adult females in the Myzolecaniinae can be differentiated from species in the other nine subfamilies known in the Coccidae by the following features: (1) lack of dorsal tubular ducts and eyespots; (2) presence of anal plates with numerous setae on the dorsal surface; (3) particularly large spiracles, with broad bands of spiracular pores between the margin and spiracles; (4) ventral tubular ducts of one type, frequently restricted to each side of the genital opening; (5) bands of (often rather spinose) setae replacing the normal pairs of long prevulvar setae; (6) reduced legs with fine claw digitules; (7) antennae of reduced size and often with fewer segments; and (8) a short anal tube (Hodgson, 1994). Currently, the Myzolecaniinae includes 20 genera, i.e., Akermes Cockerell, Alecanium Morrison, Alecanopsis Cockerell, Aztecalecanium Kondo & Williams, Bombacoccus Kondo, Cribrolecanium Green, Cryptostigma Ferris, Cyclolecanium Morrison, Foldilecanium Kondo, Halococcus Takahashi, Houardia Marchal, Megasaissetia Cockerell, Myzolecanium Beccari, Neolecanium Parrott, Octolecanium Kondo, Paractenochiton Takahashi, Pseudophilippia Cockerell, Richardiella Matile-Ferrero & Le Ruyet, Toumeyella Cockerell and Torarchus Gullan & Stewart (Gullan & Stewart, 1996; Kondo 2010a, 2011; Kondo & Williams 2004a, 2009; Kondo et al., 2005).

Steinweden (1929) postulated that New World species with reduced limbs formed a natural group, which he called the Toumeyella-group, in which he included the genera Toumeyella, Neolecanium and Pseudophilippia. Steinweden (1929) chiefly defined the Toumeyella-group by the following features: (1) legs and antennae greatly reduced, but retaining their parts; (2) marginal and stigmatic setae small and inconspicuous or absent; (3) anal region with hypopygial setae; and (4) presence of quinquelocular pores in the perivulvar region. Currently, the Toumeyella-group is composed of Akermes, Bombacoccus Cyclolecanium, Megasaissetia, Neolecanium, Octolecanium, Pseudophilippia and Toumeyella (Kondo 2010a; Kondo & Williams 2002; Kondo et al., 2005). It should be noted that Neolecanium is considered a synonym of Toumeyella by Williams & Kondo (2004b, 2009). It appears that soft scales with reduced limbs have evolved multiple times; based on the morphology of first-instar nymphs, Kondo and Williams (2002) suggested that the Myzolecaniinae is composed of several unrelated lineages. This hypothesis also is supported by a Bayesian phylogenetic analysis of the family Coccidae using DNA sequences based on 18S, 28S and partial CO1 data (T. Kondo & L.G. Cook, unpublished data).

The genus Toumeyella has a wide distribution in the New World with most species being described from the USA, suggesting that many more new species should be found in other countries in the Nearctic and Neotropical Regions (Kondo & Pellizzari, 2011). Toumeyella coffeae is the only species in the genus Toumeyella known to occur on the roots and root crown of its host, as all other known species are known from the aerial parts of their hosts. Morphologically, it differs from all its congeners by: (i) the small size of the perivulvar pores that are about the same size as the spiracular pores, (ii) the complete fusion of all segments of the legs except for the claw, and (iii) the poor segmentation of the antennae. The latter two are common features in the closely related genus Akermes Cockerell (T. Kondo, personal observation). Very few other coccids are known from the root system of their hosts. In the Myzolecaniinae, there are a couple of species that feed on the roots, i.e., Cryptostigma silveirai (Hempel), a species which appears to be restricted to the underground roots of grape vines and is an important pest of vines in Brazil (Hempel, 1900, Le Page & Piza, 1914), and C. rhizophilum Kondo from Colombia, Costa Rica, Ecuador and Panama is known from the root system of various hosts including Anthurium sp. (Araceae), Elaeis guineensis (Arecaceae), Ananas sp. (Bromeliaceae), Musa sapientum, M. textilis (Musaceae), Peristeria elata (Orchidaceae), Theobroma sp. (Sterculiaceae) and Zingiber officinale (Zingiberaceae) (Kondo, 2010b). The present paper should provide useful information for researchers working on coffee pests as well as quarantine personnel.


The author thanks Andrea Amalia Ramos Portilla (ICA) for providing the Venezuelan specimens of T. coffeae. Many thanks to Clemencia Villegas, Fabio Alonso Jáuregui, Manuel Alexis Leal G., María Fernanda Díaz Niño and Radamés Urtiaga for collecting the material studied. Special thanks to Clemencia Villegas and María Fernanda Díaz Niño for providing the photos in Figure 1. Many thanks to Dr. Nate Hardy (Auburn University, Alabama, USA) and Dr. Penny J. Gullan (The Australian National University, Canberra, Australia) for kindly reviewing the manuscript, and to Mrs. Nereida Mestre- Novoa for providing information on the Toumeyella species recorded from coffee in Cuba.



Abrahao J, Mamprim O. 1958. Cochonilha da raiz do cafeeiro. O Biológico. Sao Paulo 24: 268-271.         [ Links ]

Adenuga AO, Adeboyeku K. 1987. Notes on distribution of ant- Homoptera interaction on selected crop plants. Insect Science and its Application 8: 239-243.         [ Links ]

Aitken-Soux P. 1985. Quelques pestes et maladies généralement rencontrées dans les pépinières de café. Feuille d'Extension, Institut Interaméricain de Coopération pour l'Agriculture, Haïti 53: 1-5.         [ Links ]

Ali SM. 1968. Description of a new and records of some known coccids (Homoptera) from Bihar, India. Oriental Insects. New Delhi 1: 29-43.         [ Links ]

Ali SM. 1971. A catalogue of the Oriental Coccoidea (Part V) (Insecta: Homoptera: Coccoidea) (with an index). Indian Museum Bulletin 6: 7-82.         [ Links ]

Almeida de DM. 1973. Coccoidea de Angola. 1- Revisao das especies conhecidas. Boletim do Instituto de Investigacao Cientifica de Angola, Luanda 10: 1-23.         [ Links ]

Avasthi RK, Shafee SA. 1991. Revision of the genus Coccus Linn., in India (Insecta, Homoptera, Coccidae). Journal of the Bombay Natural History Society 88(3):329-348.         [ Links ]

Ballou CH. 1926. Los cóccidos de Cuba y sus plantas hospederas. Boletín Estación Experimental Agronómica. Santiago de Las Vegas, Cuba 51: 1-47.         [ Links ]

Barrera JF. 2008. Coffee pests and their management. Pp. 961-998, In: J.L. Capinera (ed.), Encyclopedia of Entomology. 2nd ed. Springer. Dordrecht, The Netherlands. 4346 p.         [ Links ]

Ben-Dov Y. 1970. The wax scales of the genus Ceroplastes Gray (Homoptera: Coccidae) and their parasites in Israel. Israel Journal of Entomology 5: 83-92.         [ Links ]

Ben-Dov Y. 1979. A taxonomic study of the soft-scale genus Kilifia (Coccidae). Systematic Entomology 4: 311-324.         [ Links ]

Ben-Dov Y. 1993. A systematic catalogue of the soft scale insects of the world (Homoptera: Coccoidea: Coccidae). Sandhill Crane Press, Gainesville, FL, USA. 536 p.         [ Links ]

Boboye SO. 1971. Scale insects on citrus and their distribution in Western Nigeria. Journal of Economic Entomology 64: 307-309.         [ Links ]

Carnero Hernández A, Pérez Guerra G. 1986. Coccidos (Homoptera: Coccoidea) de las Islas Canarias. Comunicaciones del Instituto Nacional de Investigaciones Agrarias, Serie: Protección Vegetal. Madrid 25: 1-85.         [ Links ]

Chazeau J. 1981. Données sur la biologie de Coelophora quadrivittata (Col.: Coccinellidae), predateur de Coccus viridis (Hom.: Coccidae) en Nouvelle-Calédonie. Entomophaga 26: 301-311.         [ Links ]

Culik MP, Martins DS, Ventura JA, Peronti ALBG, Gullan PJ, Kondo T. 2007. Coccidae, Pseudococcidae, Ortheziidae, and Monophlebidae (Hemiptera: Coccoidea) of Espírito Santo, Brazil. Biota Neotropica 7(3):1-5.         [ Links ]

Danzig EM, Konstantinova GM. 1990. [On coccid (Homoptera, Coccinea) fauna of Vietnam]. (In Russian). Trudy Zoologicheskogo Instituta Akademiya Nauk SSSR. Leningrad 209: 38-52.         [ Links ]

De Lotto G. 1955. Three new coccids (Hemipt.: Coccoidea) attacking coffee in East Africa. Bulletin of Entomological Research 46: 267-273.         [ Links ]

De Lotto G. 1956. The identity of some East African species of Saissetia (Homoptera, Coccidae). Bulletin of Entomological Research 47: 239-249.         [ Links ]

De Lotto G. 1957. On some Ethiopian species of the genus Coccus (Homoptera: Coccoidea: Coccidae). Journal of the Entomological Society of Southern Africa 20: 295-314.         [ Links ]

De Lotto G. 1958. New soft scales (Homoptera: Coccoidea: Coccidae) from Africa. Proceedings of the Royal Entomological Society of London (B) 27: 165-172.         [ Links ]

De Lotto G. 1960. The green scales of coffee in Africa south of the Sahara (Homoptera, Coccidae). Bulletin of Entomological Research 51: 389-403.         [ Links ]

De Lotto G. 1963. New species and a new genus of hard scales from East Africa (Homoptera: Coccidae). Proceedings of the Royal Entomological Society of London (B) 32: 191-200.         [ Links ]

De Lotto G. 1965. On some Coccidae (Homoptera) chiefly from Africa. Bulletin of the British Museum (Natural History) Entomology 16: 175-239.         [ Links ]

De Lotto G. 1967. A contribution to the knowledge of the African Coccoidea (Homoptera). Journal of the Entomological Society of Southern Africa 29: 109-120.         [ Links ]

De Lotto G. 1968. Second contribution to the knowledge of the African Coccoidea (Homoptera). Journal of the Entomological Society of Southern Africa 31: 83-86.         [ Links ]

De Lotto G. 1969. On a few old and new soft scales and mealybugs (Homoptera: Coccoidea). Journal of the Entomological Society of Southern Africa 32: 413-422.         [ Links ]

Fernández A. 1957. Las "escamas palomillas" del cafeto en Venezuela y avances logrados en su control químico. Agronomía Tropical (Venezuela) 7(1): 7-17.         [ Links ]

Fonseca JP da. 1957. Três novas espécies de coccídeos do Brasil, sobre cafeeiro (Homoptera - Coccidae). Arquivos do Instituto Biológico, São Paulo 24: 123-135.         [ Links ]

Gill RJ. 1988. The Scale Insects of California: Part 1. The Soft Scales (Homoptera: Coccoidea: Coccidae). California Department of Food & Agriculture, Sacramento, CA, USA. 132 p.         [ Links ]

González-Hernández H, Atkinson TH. 1984. Cocoideos (Homoptera: Coccoidea) asociados a árboles frutales de la región central de México. Agrociencia 57: 207-225.         [ Links ]

Granara de Willink MC, Pirovani VD, Ferreira PS. 2010. Las especies de Coccus que afectan Coffea arabica en Brasil (Coccoidea: Coccidae) y Redescripción de dos especies. Neotropical Entomology 39(3): 391-399.         [ Links ]

Green EE. 1889. Descriptions of two new species of Lecanium from Ceylon. Entomologist's Monthly Magazine 25: 248-250.         [ Links ]

Green EE. 1896. Catalogue of Coccidae collected in Ceylon. Indian Museum Notes 4: 2-10.         [ Links ]

Green EE. 1903. Remarks on Indian scale insects (Coccidae). With descriptions of new species. Pt. II. Indian Museum Notes 5: 93- 103.         [ Links ]

Green EE. 1904a. The Coccidae of Ceylon. Pt. III. Dulau & Co., London. 171-249.         [ Links ]

Green EE. 1904b. On some Javanese Coccidae: with descriptions of new species. Entomologist's Monthly Magazine 40: 204-210.         [ Links ]

Green EE. 1907. XII. Notes on the Coccidae collected by the Percy Sladen Trust Expedition to the Indian Ocean: supplemented by a collection received from Mr. R. Dupont, Director of Agriculture, Seychelles. Transactions of the Linnean Society of London, Zoology 12: 197-207.         [ Links ]

Green EE. 1909. The Coccidae of Ceylon. Part IV. Dulau & Co., London. 250-344.         [ Links ]

Green EE. 1916. Report on some Coccidae from Zanzibar, collected by Dr. W.M. Aders. Bulletin of Entomological Research 6: 375-376.         [ Links ]

Green EE. 1937. An annotated list of the Coccidae of Ceylon, with emendations and additions to date. Ceylon Journal of Science Section B. Zoology and Geology 20: 277-341.         [ Links ]

Gullan PJ, Stewart AC. 1996. A new genus and species of ant-associated coccid (Hemiptera: Coccidae: Myzolecaniinae) from Canthium Lam. (Rubiaceae). Memoirs of the Queensland Museum 39: 307-14.         [ Links ]

Hall WJ. 1924. The insect pests of citrus trees in Egypt. Bulletin, Ministry of Agriculture, Egypt, Technical and Scientific Service 45: 1-29.         [ Links ]

Hamon AB, Williams ML. 1984. The soft scale insects of Florida (Homoptera: Coccoidea: Coccidae). In: Arthropods of Florida and Neighboring Land Areas. Florida Department of Agriculture & Consumer Service, Division of Plant Industries, Gainesville, FL, USA. 194 p.         [ Links ]

Hanford L. 1974. The African scale insect genus Udinia De Lotto (Coccidae). Transactions of the Royal Entomological Society of London 126: 1-40.         [ Links ]

Hempel A. 1900. Descriptions of three new species of Coccidae from Brazil. Canadian Entomologist 32: 3-7.         [ Links ]

Hempel A. 1921. Três novos coccídeos. Archivos da Escola Superior de Agricultura e Medicina Veterinária. Rio de Janeiro 5: 143-146.         [ Links ]

Hodgson CJ. 1967. Notes on Rhodesian Coccidae (Homoptera: Coccoidea): Part 1: The genera Coccus, Parasaissetia, Saissetia and a new genus Mashona. Arnoldia (Rhodesia) 3: 1-22.         [ Links ]

Hodgson CJ. 1969a. Notes on Rhodesian Coccidae (Homoptera: Coccoidea): Part II: The genera Ceroplastes and Gascardia. Arnoldia (Rhodesia) 4: 1-43.         [ Links ]

Hodgson CJ. 1969b. Notes on Rhodesian Coccidae (Homoptera: Coccoidea): Part III. Arnoldia (Rhodesia) 4: 1-42.         [ Links ]

Hodgson CJ. 1994. The scale insect family Coccidae: an identification manual to genera. CAB International, Wallingford, Oxon, UK. 639 p.         [ Links ]

Ihering H von. 1897. Os piolhos vegetaes (Phytophthires) do Brazil. Revista do Museu Paulista. São Paulo 2: 385-420.         [ Links ]

Kirkaldy G. 1902. Hemiptera. Fauna Hawaiiensis 3: 93-174.         [ Links ]

Kondo T, Muñoz JA, López R, Reyes J, Monsalve J, Mesa NC. 2011. Insectos escama y ácaros comunes del aguacate en el Eje Cafetero y el Valle del Cauca, Colombia. Corpoica. Produmedios, Bogotá. 20 p.         [ Links ]

Kondo T, Pellizzari G. 2011. Description of a new species of Toumeyella Cockerell (Hemiptera: Coccidae) from Mexico; with a taxonomic key to Mexican species. Revista Brasileira de Entomologia 55 (2):229-233.         [ Links ]

Kondo T, Williams DJ. 2008. Neotype designation and redescription of Toumeyella liriodendri (Gmelin) (Hemiptera: Coccoidea: Coccidae). Journal of Insect Science 8 (56):1-6.         [ Links ]

Kondo T, Williams ML, Gullan PJ. 2005. Redescription of Octolecanium perconvexum (Cockerell), New Genus and New Combination, with Description of a New Species from Guatemala (Hemiptera: Coccoidea: Coccidae). TIP Revista Especializada en Ciencias Químico-Biológicas 8(1):5-11.         [ Links ]

Kondo T, Williams ML. 2004a. Redescription of the myrmecophilous soft scale insect: Aztecalecanium colimae (Cockerell), new genus and new combination (Hemiptera: Coccoidea: Coccidae). TIP Revista Especializada en Ciencias Químico-Biológicas 7 (1):5-9.         [ Links ]

Kondo T, Williams ML. 2004b. Redescription of the Mexican soft scale insect Toumeyella sallei (Signoret), comb.n. (Insecta: Hemiptera: Coccidae). Ann. Naturhist. Mus. Wien. B 105: 211- 215.         [ Links ]

Kondo T, Williams ML. 2009. Redescriptions of Neolecanium leucaenae Ckll., Toumeyella cerifera Ferris and T. sonorensis Ckll. & Parrott and their transfer to Neotoumeyella gen. nov. (Hemiptera: Coccidae), with descriptions of two new species from the southeastern USA and Colombia, South America. International Journal of Insect Science 2: 11-27.         [ Links ]

Kondo T, Williams ML. 2002. The Myzolecaniinae (Hemiptera: Coccidae): Old World vs. New World. Bolletino di Zoologia Agraria e di Bachicoltura, Ser. II, 33 (3):125-128.         [ Links ]

Kondo T. 2006. A new African soft scale genus, Pseudocribrolecanium gen. nov. (Hemiptera: Coccoidea: Coccidae), erected for two species, including the citrus pest P. andersoni (Newstead) comb. nov. Journal of Insect Science 6(1):1-16.         [ Links ]

Kondo T. 2010a. Description of a new coccid (Hemiptera, Coccidae) on avocado (Persea americana Mill.) from Colombia, South America. Zoo Keys 42: 37-45.         [ Links ]

Kondo T. 2010b. Taxonomic revision of the myrmecophilous, meliponiphilous and rhizophilous soft scale genus Cryptostigma Ferris (Hemiptera: Coccoidea: Coccidae). Zootaxa 2709: 1-72.         [ Links ]

Kondo T. 2011. Transfer of the myrmecophilous soft scale insect Neolecanium amazonensis Foldi to Foldilecanium gen. nov. (Hemiptera: Coccidae), with description of a new species from Colombia. Insecta Mundi 0167: 1-10.         [ Links ]

Laing F. 1929. Descriptions of new, and some notes on old species of Coccidae. Annals and Magazine of Natural History 4: 465-501.         [ Links ]

Le Pelley RH. 1968. Pests of coffee. Longmans, London. 588 p.         [ Links ]

Lepage HS, Piza MT. 1941. Redescrição do Neolecanium silveirai (Hempel) (Homoptera-Coccoidea), séria praga da videira e seu controle. Arquivos do Instituto Biológico. São Paulo 12: 21-26.         [ Links ]

Lit IL. 1997. First report of the family Lecanodiaspididae and other new records and notes on Philippine scale insects (Coccoidea, Hemiptera). Philippine Entomologist 11: 87-95.         [ Links ]

Mamet JR. 1959. Notes on the Coccoidea of Madagascar IV. Mémoires de l'Institut Scientifique de Madagascar (Ser. E) 11: 369-479.         [ Links ]

Mestre Novoa N, Hamon A, Evans G, Kondo T, Oliver P, Marrero A, Alonso A. 2011. Los cocoideos (Hemiptera: Sternorrhyncha: Coccoidea) presentes en la Cordillera de Guaniguanico, Pinar del Río, Cuba, y la relación con sus hospedantes. Insecta Mundi 0183: 1-25.         [ Links ]

Murphy ST. 1991. Insect natural enemies of coffee green scales (Hemiptera: Coccidae) in Kenya and their potential for biological control of Coccus celatus and Coccus viridis in Papua New Guinea. Entomophaga 36: 519-529.         [ Links ]

Murphy ST. 1997. 3.3.14 Coffee. 367-380. In: Ben-Dov Y. & Hodgson CJ (Editors), Soft Scale Insects - Their Biology, Natural Enemies and Control [Vol. 7B]. Elsevier, Amsterdam & New York. 442 p.         [ Links ]

Muzaffar N, Ahmad R. 1977. A note on Saissetia privigna (Hem.: Coccidae) in Pakistan and the breeding of its natural enemies. Entomophaga 22: 45-46.         [ Links ]

Nakahara S, Miller CE. 1981. A list of the Coccoidea species (Homoptera) of Puerto Rico. Proceedings of the Entomological Society of Washington 83: 28-39.         [ Links ]

Nakahara S. 1981. List of the Hawaiian Coccoidea (Homoptera: Sternorhyncha). Proceedings of the Hawaiian Entomological Society 23: 387-424.         [ Links ]

Newstead R. 1911. On a collection of Coccidae and Aleurodidae, chiefly African, in the collection of the Berlin Zoological Museum. Mitteilungen aus dem Zoologischen Museum in Berlin 5: 155-174.         [ Links ]

Newstead R. 1917. Observations on scale-insects (Coccidae) - V. Bulletin of Entomological Research 8: 125-134.         [ Links ]

Newstead R. 1920. Observations on scale-insects (Coccidae) - VI. Bulletin of Entomological Research 10: 175-207.         [ Links ]

Ramakrishna Ayyar TV. 1919. A contribution to our knowledge of South Indian Coccidae. Bulletin of the Agricultural Research Institute, Pusa, India 87: 1-50.         [ Links ]

Ramakrishna Ayyar TV. 1930. A contribution to our knowledge of South Indian Coccidae (Scales and Mealybugs). Bulletin of the Imperial Institute of Agricultural Research, Pusa, India 197: 1-73.         [ Links ]

Shafee SA, Yousuf M, Khan MY. 1989. Host plants and distribution of coccid pests (Homoptera: Coccoidea) in India. Indian Journal of Systematic Entomology 6: 47-55.         [ Links ]

Sheffer BJ, Williams ML. 1990. Descriptions, distribution, and host-plant records of eight first instars in the genus Toumeyella (Homoptera: Coccidae). Proceedings of the Entomological Society of Washington 92(1):44-57.         [ Links ]

Silva d'Araujo GA, Goncalves CR, Galvao GM, Goncalves DM. 1968. Fourth catalog of insects that live in Brazil. Part II. Insects, hosts and natural enemies. Ministerio da Cultura, Rio de Janeiro. V. 1, 622 p.         [ Links ]

Steinweden JB. 1929. Bases for the generic classification of the coccoid family Coccidae. Annals of the Entomological Society of America 22: 197-245.         [ Links ]

Vayssière P. 1955. Les animaux parasites du caféier. (In French). p. 257-267. In: Coste R. (Editor), Les Caféiers et les Café dans le Monde, Paris. 382 p.         [ Links ]

Williams DJ, Watson GW. 1990. The Scale Insects of the Tropical South Pacific Region. Part 3. The soft scales (Coccidae) and other families. CAB International, Wallingford, UK. 267 p.         [ Links ]

Williams DJ. 1982. The distribution and synonymy of Coccus celatus De Lotto (Hemiptera: Coccidae) and its importance on coffee in Papua New Guinea. Bulletin of Entomological Research 72: 107-109.         [ Links ]

Williams DJ. 1986. Scale insects (Homoptera: Coccoidea) on coffee in Papua New Guinea. Papua New Guinea Journal of Agriculture, Forestry and Fisheries 34: 1-7.         [ Links ]

Williams ML, Kondo T. 2001. Characteristics of first-instar nymphs in the soft scale insects (Hemiptera: Coccidae): surprising indicators of relationships. Bollettino di Zoologia Agraria e di Bachicoltura (Milano) 33(3): 35-42.         [ Links ]

Williams ML, Kondo T. 2008. Status and current composition of the soft scale insect genus Toumeyella (Hemiptera: Coccoidea). Proceedings of the XIth International Symposium on Scale Insect Studies. Editors: Branco M, Franco JC & Hodgson C. Oeiras, Portugal. September 24-27, 2007. Pp. 29-32.         [ Links ]

Williams ML. 1993. Toumeyella lignumvitae, a new species of scale insect from the Florida keys (Homoptera: Coccidae). Florida Entomologist 76 (4):566-572.         [ Links ]

Williams ML. 2010. Annotated list of the scale insects of Guatemala. Entomologia Hellenica 19: 144-152.         [ Links ]

Wolcott GN. 1941. Supplement to Insectae borinquenses. Journal of Agriculture of the University of Puerto Rico 25: 33-158.         [ Links ]