Corpoica. Ciencia y Tecnología Agorpecuaria ISSN: 0122-8706 [email protected] Corporación Colombiana de Investigación Agropecuaria Colombia

Kondo, Takumasa A new species of Toumeyella Cockerell (: Coccoidea: ) on roots, arabica L. (Rubiaceae), from Colombia and Venezuela Corpoica. Ciencia y Tecnología Agorpecuaria, vol. 14, núm. 1, enero-junio, 2013, pp. 39- 51 Corporación Colombiana de Investigación Agropecuaria Cundinamarca, Colombia

Available in: http://www.redalyc.org/articulo.oa?id=449945181006

How to cite Complete issue Scientific Information System More information about this article Network of Scientific Journals from Latin America, the Caribbean, Spain and Portugal Journal's homepage in redalyc.org Non-profit academic project, developed under the open access initiative Revista Corpoica - Ciencia y Tecnología Agropecuaria enero - junio / 2013 p. 39-51 39

GENÉTICA VEGETAL Y BIODIVERSIDAD

ARTÍCULO CIENTÍFICO

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 Corporación Colombiana de Investigación Agropecuaria (Corpoica), Centro de Investigación Palmira, Calle 23, Carrera 37, Palmira, Valle, Colombia, [email protected]

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

ABSTRACT RESUMEN

A new species of soft scale from Colombia and Una nueva especie de escama blanda de Colombia y Venezuela, Toumeyella coffeae Kondo sp. nov., is described Venezuela, Toumeyella coffeae Kondo sp. nov., se describe and illustrated based on the adult female. A taxonomic key e ilustra con base en la hembra adulta. Se proveen una to the genera of scale of the Toumeyella-group clave taxonómica para separar los géneros de escamas and a list of 49 soft (Coccidae) species recorded blandas del grupo-Toumeyella y una lista de 49 especies from Coffea spp. worldwide are provided. de escamas blandas (Coccidae) registradas sobre Coffeaspp., en el mundo.

Key words: coccid, scale insect, soft scales,Toumeyella Palabras claves: cóccido, insecto escama, escama coffeae. blanda, Toumeyella coffeae.

© 2013 Corporación Colombiana de Investigación Agropecuaria GENÉTICA VEGETAL p. 39-51

Y BIODIVERSIDAD 40 A new species of Toumeyella Cockerell (Hemiptera: Coccoidea: Coccidae) on coffee roots,Coffea arabica L. (Rubiaceae), from Colombia and Venezuela enero - junio / 2013 1968; Murphy, here described 1997).The fromspecies 2008;Fernández, Pelley, 1957;Le (Barrera, host its of Venezuela, where itaffects the and roots crownroot for alongconsidered in time ofcoffee apest as been known in the literature as and scale soft of species named 15 insects in included the genus are there Currently, described (2011) & Parrott), andmore recently Kondo &Pellizzari Neotoumeyella cerifera cerifera species. some Neotoumeyella of morphology the Later, Kondo &Williams the (2009)erected genus on comments and information about the distribution, host plants scale insect pests. scale to quarantineaid personnel and researchers working on on reported Coccidae) insects (Hemiptera: Coccoidea: scale Central and South America. Furthermore, alist of soft identificationeasy genera ofclosely related thatoccur in of (excluding separate the in genera included the Toumeyella-group species andillustrated. isdescribed Ataxonomic key to of of species new The purposemain of paper this is the description ofa personal communication). hereinthe species (N.Mestre-Novoa, new described isdifferent fromof the plant andthe morphology in on coffee reported comes fromCuba the aerial parts confirmation.this needs On the other hand, the species in ofcoffee apest aboveas reported however,Venezuela, from roots also coffee maycorrespond to the species specimens from andVenezuela, Colombia collected There is another of undescribed species Santana andTrinidad cultivars (Kondo americana al sylvestris Cuba on Therefurther are from recorded undescribed species the United States (9spp.) (Kondo &Pellizzari, 2011). in (2spp.), Brazil Cuba (1sp.), Mexico (5spp.) and in thespecies genus of16 Williams andKondo anoverview (2008)gave INTRODUCTION ., 2011)andfrom on Colombia , Coffea arabica Toumeyella Coffea Eugenia foetida Eugenia Ferris and Sw. (Salicaceae), Adelia ricinella spp. worldwide (Rubiaceae) an is provided as L., on 8,Choquette, 7,Booth Booth L., Lorena, Neolecanium (Kondo, 2004b, 2009) is provided for into transferred they which Toumeyella plants. Theadult female of new this Toumeyella fontanai T. sonorensis Pers. (Myrtaceae) (Mestre (Ferris) and Toumeyella that is considered a synonym L. (Euphorbiaceae), L. Coffea arabica collected from the roots collected Toumeyella Toumeyella Cockerell, providing Ckll. &Parrott as N. sonorensis from Mexico. et al et (Rubiaceae) (Rubiaceae) sp. that has has that sp. , distributed Toumeyella Toumeyella Casearia ., 2011). (Ckll. Persea et et , Universidad Nacional Bogotá, sede deColombia, UNAB: in the isdeposited studied following institutions: of important features around The thematerial margin. side andthe venter on the sidewith right enlargements the adult female 2) shows (Fig. the dorsum on the left specimens in the material Theillustration studied. of determined by selecting the smallest andthe largest are in given parenthesis. size body was of The range Measurementsincluding the holotype). ofthe holotype mounted specimens for eachofthe fivecollecting sites, wereand taken from 10specimens (the slide- two best Pellizzari& (2011). Measurements a range are as given mostly that ofKondo &Williams (2008) and Kondo features follows ofmorphological The terminology bydescribed Williams &Granara (1992). deWillink Specimens were slide-mounted following the procedure – Anal plates nearmid-dorsum, notlocated usually 1. 2009): Kondo (adapted &Williams, from Kondo,females 2010a; toKey genera ofthe Toumeyella-group on adult based RESULTS DISCUSSION AND name in was checked ScaleNet. ofeachspecies additional information from the literature. The valid insect database ScaleNet (Ben-Dov mostlycompiled from information found in the scale on reported Coccidae) insectsA listofsoftscale (Hemiptera: Coccoidea: Cali, Valle delCauca, Colombia. UVCO: Cundinamarca,Bogotá, Colombia. MATERIALS AND METHODS AND MATERIALS

marginal setaemarginal variable, ……...... ……..2 notpeg-like complete, both extending pore bands to margin; anteriormargin; and posterior spiracular pore bands from atlength posterior about 1/5to 1/4ofbody found ……………………………...….…...... peg-like setaemargin); marginal cylindrical, spiracular (posterior extending pore band margin to incomplete, pore band spiracular notextending to Anal plates anterior nearmid-dorsum; located Museo Facultad Entomológico deAgronomía, Museo Universidad de Entomología, del Valle, Revista Corpoica -Ciencia y Tecnología Agropecuaria Coffea Cyclolecanium spp. worldwide was et al et ., 2013)with Morrison

© 2013Corporación Colombiana deInvestigación Agropecuaria stigmatic clefts shallow or absent ...... and sharply setae spinose, generally rarely lanceolate; vulva around extending dorsal anteriorly line; in amedio-lateral located ducts tubular Ventral – 7. – Ventral tubular ducts notdistributed above………… as 6. Dorsum ofslide-mounted specimens without – adense 5. Ventral – tubular ducts present at around least perivulvar 4. variable, microducts dorsal present; pores Preopercular – 3. microducts around notconspi– Dorsal margin body 2.

often also on posterioroftenalso on segments, abdominal but not matic clefts deep……………….. stig segment setaeabdominal V; dorsal lanceolate; tending anteriorly line up to in about a medio-lateral …………...... …………………... 7 ...... posterior spiracular ...... ………. pore band around andreachingarea on region band abdominal ……………………...... …….. with andwith 7or 2or 8loculi 3central ...... loculi Perivulvarby athin wax. layer pores mostly ofglassy pattern ofmicroducts. Adult females incovered life …………………………………...……… pores mostly and one with central 5 loculi loculus in disappears specimens. old Perivulvar wax powdery life this in females adult covered although by awhite cottony wax, to powdery Young microducts. of pattern area ………………...... ……….6 invaginated of …...... 4 pattern dense stigmatic microducts; spinesbilocular usually present without generally absent ………….……...... … stigmaticof invaginated microducts; bilocular spines …………………………………………....……………………….. 3 thancuously larger rest ofmicroducts on dorsum ………………………………………….. thancuously larger rest ofmicroducts on dorsum …………………………………...... ….. Ventral tubular ducts around located vulva and ex Ventral tubular ducts present at in least asubmarginal Dorsum ofslide-mounted specimens with adense Ventral tubular ducts absent……………………………… 5 Preopercular pores absent; dorsum with dense pattern microducts around conspi-Dorsal margin body Neotoumeyella Pseudophilippia Megasaissetia Toumeyella Kondo andWilliams Octolecanium Bombacoccus Akermes Cockerell Cockerell Cockerell Cockerell Kondo Kondo - - - 8 first-instar8 nymphs)(UNAB). Municipio Arboledas, Niño,Díaz excafé Canastilla, 20(26adult females + °W, 10.iv.2013, 1,497ma.s.l., María coll. Fernanda Cucutilla, 07,34513 Finca°N, 072,47227 La Dalia, + 3female third-instar nymphs) Municipio (UNAB). second-instar nymphs + 4female second-instar nymphs 36 (28adult females +28first-instar nymphs +8 male Fabiocoll. Alonso Jáuregui, exroots of Finca vereda Aguadas, ElArboledas, Porvenir, 2.v.2013, females +3third-instar nymphs) Municipio (UNAB). Paratypes. Manuel ex rootscoll. of Alexis Leal, Vereda Finca Bajo, Pedregal Buenos Aires, 2.v.2013, Norte deSantander: municipio deCucutilla, Colombia: Material studied. Escama blandaSpanish: delcafé. dela raíz scale. Coffee-root Proposed English: common name: Toumeyella coffeae scattered. Simple pores (sp) eachabout 2.2–3.0 µm µmwide, with along terminal2.0–2.5 filament,evenly less scattered evenly. microducts (dmic) Dorsal each rather scarse, moreeach 8.6–20.0(9–17)µmlong, or setae (dset) sharply spinose, straight curved, or slightly heavilybecoming sclerotized in older specimens. Dorsal Dorsum. 1.1–4.1(1.4)mm wide (Figuremm 2). long, oftenoval, narrowing 1.7–5.3(1.8) body anteriorly; Mounted material. 4.2 mm in diameter (Figure 1). anal plates dark brown. to reddish Mature insects 1.5– but usually with mid-dorsum or notmottled; lightly very brown in color, green to greyish with dark tesselations, Unmounted material. Description. Adult female (Figures 1&2) female third-instar nymph) (UNAB). first-instar nymph +1female second-instar nymph +1 Urtiaga, #1255,2(2adult females Radamés +1 coll. vi.2010, Sanare, Arriba, 1,100ma.s.l., Sabana Redonda Other material studied. (UVCO). 17 first-instar nymphs +1female third-instar nymph) Coffea arabica G.,exroots of Clemencia coll. 25.xi.2012, Villegas vereda Finca LaRinconada, El 1,526ma.s.l., Cedro, Derm membranous on young adult females, Same data as holotype 31 slides (28adult 31slides Same data holotype as var. 20(15adult females + Colombia,

Holotype. Kondo, sp. nov. (Figures 1&2)

Body outlineBody oval to elongate Body convex. Derm Body orange,

Venezuela: Estado deLara:

Adult female (UNAB). Coffea arabica Coffea arabica , .

GENÉTICA VEGETAL p. 39-51

Y BIODIVERSIDAD 41 A new species of Toumeyella Cockerell (Hemiptera: Coccoidea: Coccidae) on coffee roots,Coffea arabica L. (Rubiaceae), from Colombia and Venezuela enero - junio / 2013 GENÉTICA VEGETAL p. 39-51

Y BIODIVERSIDAD 42 A new species of Toumeyella Cockerell (Hemiptera: Coccoidea: Coccidae) on coffee roots,Coffea arabica L. (Rubiaceae), from Colombia and Venezuela enero - junio / 2013 (13–36) µm long. Eyes(13–36) µmlong. notdetected. lateral spines 15–22 29–44(27–42) µmlong, longest, to sharply stigmatic spinose, median spine usually spines stigmatic per stigmatic spines area; (stgsp) bluntly Stigmaticareas. usually with 3 clefts developed, well anterioron eachsidebetween and posterior stigmatic often row,a single, irregular with 5–10(6 on holotype) in arranged bent, slightly to each12–50 (17–30) µm long, Margin. area often present just anterior plates.to anal setae (setae represented by dark dots). Asmall sclerotic setae. Analsetae ofhypopygial ring and 5 pairs with 10 oflongplus setae, 1pair fringe about 7ventral subapical with 1or 2setae surface, on dorsal (140–142) µmlong, posterolateral 113–147 margin (125–142) µmlong, 118 (98–110)µmwide, anterolateral 123–164 margin 93– eachplatemargin, 167–203(183–191) µmlong, fromat length posterior about 1/5to 1/4ofbody Anal plates (aplt) quadrate, together plates located tuberclesDorsal andpocket-like sclerotizations absent. just dorsad to labium, eachpore 5–14(6–11)µmwide. anterior to anal plates, extending anteriorly up to area Preopercular present pores (prop) on mid-dorsum wide, scattered. evenly tubular ducts Dorsal absent. Main photo by C. Villegas; upperandlower insetphotos by M.F. Niño. Díaz Figure 1. Toumeyella coffeae Marginal setae (mset) sharply spinose, straight Kondo onroots of Coffea arabica (Norte deSantander,(Norte Colombia). tarsal digitules knobbed, longer than longer claw knobbed, digitules. digitules tarsal without adenticle, slender, claw digitules, knobbed; clawsmetathoracic well developed, usually largest; legs including claw,legs each59–105 (80–102)µmlong, except for claw,discernible, fused, ofall total length segments not greatly reduced, (70–71) µmwide. Legs 78 (61–65)µmwide, posterior peritremes each66–88 anteriorwell developed, spiracular peritremes each56– laterally from Spiracles eachspiracle margin. to body pores wide, rarely up to 6in some sections, extending pores with present more loculi, in anarrow 1–3 band wide, mostly with rarely with 4–6loculi, afew 5 loculi, metathoracic Spiracular pores (spp) leg. each5–6 µm on segment eachabdominal andlateral tosubmedially andin small groupssegment ofabout VII, 1–6pores more,or present around on abdominal afew vulvar area, (mostly rarely up afew to withloculi 8loculi 5loculi), spiracularsame sizeas pores, each5–6µmwide, with 4–6 wide. Perivulvar rather pores (pvp) scarce, about small, duct terminal opening branched small, 4–5µm gland, each tubular duct with aterminal filament ending in a tral tubular ducts scarce, present around vulvarregion, scattered throughout, evenly eachabout 3µmwide. Ven Ventral longest. last segment (VII) microducts (vmic) on apair setae,of long each23–70µmlong, median 3pairs also straight bent, or slightly each6–27µmlong; Venter. Upper inset. Derm entirely membranous. Ventral setae slender, Young adultfemales. Revista Corpoica -Ciencia y Tecnología Agropecuaria Lower inset . Old adultfemale.. Old -

© 2013Corporación Colombiana deInvestigación Agropecuaria = simplepore; spp=spiracular pore; stgsp =stigmatic spine;vmic=ventral microduct; vset=ventral setae;vtbd =ventral tubularduct. Abbreviations: aplt=anal plate; =antenna; ar=analring; dmic=dorsalmicroduct; dset=dorsalsetae;msetmarginal setae; prop =preopercular pores; pvp =perivulvar pore; sp Figure 2. Toumeyella coffeae Kondo, adultfemale

GENÉTICA VEGETAL p. 39-51

Y BIODIVERSIDAD 43 A new species of Toumeyella Cockerell (Hemiptera: Coccoidea: Coccidae) on coffee roots,Coffea arabica L. (Rubiaceae), from Colombia and Venezuela enero - junio / 2013 GENÉTICA VEGETAL p. 39-51

Y BIODIVERSIDAD 44 A new species of Toumeyella Cockerell (Hemiptera: Coccoidea: Coccidae) on coffee roots,Coffea arabica L. (Rubiaceae), from Colombia and Venezuela enero - junio / 2013 Table 1. mealybug 2013, Norte Bajo, Pedregal deSantander, on 2.v Colombia, stems. In the from collection Finca Buenos Aires at Vereda on roots andon undergroundlive andfeed of parts males have found been in one infestation.coccids These Biology. Williams, 1996;Williams, 1993). Toumeyella antennaesegmented andresemble other of species Notes. coffee. Etymology. setae. of labial labium with 1 segmented, 4 pairs 240 (240) µm wide; 167– shield Mouthparts clypeolabral welldeveloped, ofinterantennalpairs setae, each5–32(5–20)µmlong. with fleshy setae present on ofantennae.apex With 3 segmentation 2–4 (2) segmented, developed, poorly Antennae (ant) each44–100(73–84)µmlong, short, Ceroplastes giganteus Ceroplastes galeatus Ceroplastes floridensis Ceroplastes destructor Ceroplastes brevicauda Avricus arborescens Avricus amoenus Alecanochiton marquesi Toumeyella coffeae Species ofCoccidae recorded on First-instar nymphs of Dysmicoccus brevipes This species issexualbecause second-instar This species (Kondo &Williams, 2002;Sheffer & The species is named after is The named itsspecies host: Type Species (De Lotto) (De Newstead (Laing) Dozier Newstead Comstock Hall Hempel was found with together the (Cockerell) (Pseudococcidae).(Cockerell) Coffea spp. worldwide T. coffeae Guatemala Uganda Uganda, Zimbabwe Sao Tome,Sudan, Tanzania, Angola, Ethiopia, Kenya, Ghana, Zimbabwe Brazil Brazil (Espiritu Santo), India,Israel Uganda of theCongo, ofthe Congo, Republic Angola, Cameroon, Democratic Republic Angola, Kenya, Uganda have five- have Country ofrecordCountry Gill, 1988;HamonGill, &Williams, 1984). the aerial ofits parts host (Kondo &Williams, 2008; T. liriodendri Table this 1because islikely amisidentification since the in rootsGuatemala ofcoffee from omitted was liriodendri al et country in records ScaleNet not listed (Ben-Dov on recorded has been each species here. Table the country 1gives or countries from which Coffea Dov, insects ofsoftscale on 1993).Atotal of49species Ramakrishna Ayyar, isa which identifiedgenus to only and level on recorded Dov to the insectdatabase scale ScaleNet (Ben-According spp. in the world (Nietner), thus its distribution under islisted Murphy (1997)isasynonym of Pelley by by Le (1968)andcited recorded Lindinger Soft scale insectsSoft (Coccidae) on coffee, et al et ., 2013)andthe validation source. spp., including the species, new ., 2013)there insects ofsoftscale are 47species (Gmelin)by Barrera listed (2008)on Coffea is aNorth American that on species feeds Ben-Dov Ben-Dov Le Pelley (1968);Murphy (1997) Culik (1970);Ben-Dov Ben-Dov (1997) LottoDe (1965);Le Pelley (1968);Murphy Almeida (1973);Ben-Dov Murphy (1997) Lotto (1955,1965;1967);Le Pelley (1968); Almeida (1973);Ben-Dov Murphy (1997) Lotto (1968);Laing (1929);Le Pelley (1968); Almeida (1973);Ben-Dov Hodgson (1969a) Ben-Dov Le Pelley (1968);Murphy (1997) spp. worldwide, excluding species et al Revista Corpoica -Ciencia y Tecnología Agropecuaria et al . (2007);Hodgson (1994) et al . (2013); Williams (2010) . (2013); De Lotto. (2013);De (1958); References Coffea nomen nudum Lecanium mercarae nigra T. coffeae et al et al et al asiaticus spp., additional et al . (2013); . (2013); De . (2013);De . (2013);De Toumeyella . (2013); , are listed P. nigra Coffea (Ben-

© 2013Corporación Colombiana deInvestigación Agropecuaria Coccus alpinus Coccus africanus Ceroplastes vinsonioides Ceroplastes stellifer Ceroplastes rubens Ceroplastes personatus Ceroplastes luteolus Coccus colemani Coccus celatus Coccus brasiliensis Coccus subhemisphaericus Coccus subacutus Coccus longulus Coccus lizeri Coccus hesperidum (Fonseca) De Lotto De De Lotto De (Douglas) Species Kannan (Newstead) (Newstead) Fonseca Maskell (Westwood) De LottoDe L. Newstead Newstead (Newstead) India Vietnam Philippines, Sudan, Tanzania, Uganda, Kenya,Brazil, Malaysia, Papua New Guinea, Brazil Tanzania, Uganda,Zimbabwe of theCongo, Eritrea, Ethiopia, Kenya, DemocraticAngola, Brazil Republic (Bahia), Kenya, Nigeria Nigeria, Tanzania, Uganda,Zimbabwe Kenya, Guinea-Bissau, Angola, Ghana, Ghana Western Samoa Nigeria, Sudan,Uganda,ZimbabweGhana, Kenya Angola, Ghana, Kenya,Angola, Ghana, Tanzania, Uganda Uganda Vanuatu Brazil Peru, Tanzania Angola, Guatemala, Guyana, Mexico, Country ofrecordCountry (1968); Murphy (1997);Newstead (1917) Ben-Dov Newstead (1911) (1969b); Le Pelley (1968);Murphy (1997); Lotto (1971);De (1965,1968);Hodgson Boboye Almeida (1973);Ben-Dov Le Pelley (1968);Murphy (1997) Williams & Watson (1990) Ben-Dov Newstead (1917) (2012); Le Pelley (1968); Murphy (1997); Ben-Dov Ben-Dov Ramakrishna (1919,1930);Shafee Ramakrishna et al Ali (1971);Avasthi &Shafee(1991);Ben-Dov (1968); Murphy (1997); Williams (1982) 1969); Granara de Willink Danzig &Konstantinova Lotto (1990);De (1969; Ben-Dov Murphy (1997) de Willink Ben-Dov (1967); Le Pelley (1968);Murphy (1997) de Willink Granara LottoAlmeida (1973);De (1960, 1965,1968); Pelley (1968);Murphy (1997);Newstead (1917) Ben-Dov (1968); Murphy (1997);Newstead (1920) Ben-Dov Ben-Dov (1997 de Willink Ben-Dov (1911) Le Pelley (1968);Murphy (1997);Newstead et al Almeida (1973);Barrera (2008);Ben-Dov . (2013);Le Pelley (1968);Murphy (1997); . (2013);González &Atkinson (1984); et al et al et al et al et al et al et al et al et al et al et al et al . (2013); De Lotto. (2013);De (1955,1965;1967) . (2013); Williams & Watson (1990) . (2013); De Lotto. (2013);De (1957);Le Pelley . (2013); De Lotto. (2013);De (1957);Le Pelley . (2013); De Lotto. (2013);De (1957,1967);Le . (2013); Buckley & Gullan (1991); &Gullan . (2013);Buckley . (2013);Hodgson &Peronti . (2013);Fonseca (1957);Granara . (2013);Fonseca (1957);Granara . (2010);Le Pelley (1968);Murphy . (2013);Brimblecombe (1956); . (2010);Le Pelley (1968); References et al et al et al . (2010); Hodgson . (2010);Le Pelley . (2013); et al . (1989)

GENÉTICA VEGETAL p. 39-51

Y BIODIVERSIDAD 45 A new species of Toumeyella Cockerell (Hemiptera: Coccoidea: Coccidae) on coffee roots,Coffea arabica L. (Rubiaceae), from Colombia and Venezuela enero - junio / 2013 GENÉTICA VEGETAL p. 39-51

Y BIODIVERSIDAD 46 A new species of Toumeyella Cockerell (Hemiptera: Coccoidea: Coccidae) on coffee roots,Coffea arabica L. (Rubiaceae), from Colombia and Venezuela enero - junio / 2013 Pulvinaria mammeae Pulvinaria aethiopica (Newstead) Pseudocribrolecanium andersoni Protopulvinaria longivalvata Paralecanium marianum Megapulvinaria maxima Kilifia americana Kilifia acuminata Hemilecanium imbricans Eucalymnatus tessellatus Cryptostigma inquilinum Coccus viridulus Coccus viridis (Green) De Lotto De Species (Signoret) Ben-Dov (Nietner) (De Lotto) (De Maskell (Green) (Green) (Newstead) Cockerell (Signoret) Green

Zimbabwe Lanka, Tanzania,Uganda, Vanuatu, Indies, West Papua Rico, Puerto New Guinea, Sri Reunion, Jamaica, Kenya, Malaysia, New Caledonia, of theCongo, El Salvador, Guatemala, India, Cameroon, Islands, Canary Democratic Republic Brazil Vietnam Mexico Rico Puerto India Seychelles El Salvador, Rico Puerto Kenya, Uganda Hawaii Angola, Zambia Angola Guadeloupe, Lesser Antilles Venezuela, Vietnam, WesternSamoa Taiwan, Tanzania, Tonga,Uganda, Vanuatu, Seychelles, Africa, South Sri Lanka, Surinam, Philippines, Rico, Puerto Sao Reunion, Tome, New Caledonia, Panama, Papua New Guinea, Malaysia, Mariana Islands, Mauritius, Mexico, Irian Jaya), Jamaica, Kenya, Madagascar, Hawaii, Honduras, India,Indonesia(Java, Guatemala, Guadeloupe, Guyana, Haiti, Salvador, Egypt, Ethiopia, Fiji, Ghana, the Congo, Republic, Dominican Ecuador, El Costa Cuba, Rica, Democratic of Republic Brazil,Bolivia, Cape Burma, Verde, Colombia, Country ofrecordCountry (1997); Wolcott(1941) Ben-Dov Le Pelley (1968) Ben-Dov (1902); Murphy (1997) Ben-Dov (1965); Le Pelley (1968);Murphy (1997) Almeida (1973);Ben-Dov Ben-Dov Watson (1990) Newstead (1917);Shafee Hodgson (1967);Le Pelley (1968);Murphy (1997); Carnero &Perez (1986);Green (1896,1904a,1937); Ali (1968);Barrera (2008);Ben-Dov Watson (1990) Newstead (1917);Shafee Hodgson (1967);Le Pelley (1968);Murphy (1997); Carnero &Perez (1986);Green (1896,1904a,1937); Ali (1968);Barrera (2008);Ben-Dov Le Pelley (1968);Murphy (1997) Ben-Dov (1979);Ben-Dov Ben-Dov Ben-Dov Ben-Dov Ben-Dov Barrera (2008); Ballou (1926); Ben-Dov Barrera (1926);Ben-Dov (2008);Ballou Aitken-Soux (1985);Avasthi & Shafee (1991); Squire, 1972; Williams & Watson (1990) (1959); Nakahara (1981); Shafee (1924); Ihering (1897);Le Pelley, 1968;Mamet et al Lotto 1925; Granara (1978); Ghosh, de Willink (2013); Chazeau, 1981;Culik . (2010); Green (1889, 1896, 1916); Hall Revista Corpoica -Ciencia y Tecnología Agropecuaria et al et al et al et al et al et al et al et al . (2013);Danzig &Konstantinova (1990) . (2013); De Lotto. (2013);De (1960,1969); . (2013); Brink (1992);Kondo. (2013);Brink (2006) . (2013);Nakahara (1981) &Miller . (2013);Green (1903) . (2013);Green (1907) . (2013);Le Pelley (1968); Kirkaldy . (2013);Le Pelley (1968);Murphy References et al et al et al et al . (2013) . (1989); Williams & . (1989); Williams & et al . (2013); De Lotto. (2013);De et al . (2010); De . (2010);De et al et al . (1989); . (2013); . (2013); et al .

© 2013Corporación Colombiana deInvestigación Agropecuaria (2)presence ofanal platesand eyespots; with numerous the following features: tubular (1) lack ducts ofdorsal in the other known nine subfamilies in the by Coccidae in the Myzolecaniinae differentiated can be from species cuticle Theadult females andhave appendages. reduced thatinclude species lack anumber ofstructures ofthe Myzolecaniinae (1994)to by was erected Hodgson theas Toumeyella-group (Steinweden, 1929). The well Myzolecaniinae 1994) as subfamily (Hodgson ofgenusThe soft scales the DISCUSSION privignaSaissetia oleae Saissetia neglecta Saissetia jocunda Saissetia coffeaeSaissetia Udinia paupercula Udinia glabra Udinia farquharsoni (Newstead) Udinia catori Toumeyella coffeae zanzibarensisSaissetia (Green) De Lotto De (Olivier) Maskell (Walker) Species De Lotto De De Lotto De De Lotto De De Lotto De Kondo Williams Toumeyella Sudan Uganda Democratic oftheCongo, Republic Sierra Leone Nigeria Colombia, Venezuela Tanzania India, Kenya Jamaica, Kenya, Rico Puerto Brazil, Cuba, Republic, Dominican India, Fiji, Rico, Puerto Tonga Kenya Tanzania,Uganda, Venezuela, Vietnam Seychelles, Sri Lanka, St Helena, Surinam, Papua Rico, Puerto New Guinea, Reunion, (Java), Jamaica, Kenya, Madagascar, Mexico, Guatemala, Guyana, Honduras, India, Indonesia Republic,Dominican El Salvador, Fiji, Guam, Angola, Brazil, Colombia, Costa Cuba, Rica, Tanzania, Uganda Papua Rico, Puerto New Guinea, Sri Lanka, Kenya, Mariana Islands,Jamaica, New Caledonia, Hawaii, India, Indonesia (Java, Sumatra), oftheCongo,Republic Republic, Dominican Angola, Caroline Islands, Cuba, Democratic belong to the Country ofrecordCountry & Williams, Morrison, i.e., includes 20 genera, 1994).Currently, (Hodgson, tube the Myzolecaniinae and often with anal and(8)ashort fewer segments; (7)antennaewith fine claw size digitules; of reduced legs (6)reduced normal oflong setae; pairs prevulvar of (often(5) bands rather spinose) setae replacing the frequently restricted opening; to eachsideofthe genital (4) ventraland spiracles; tubular ducts of one type, broad the ofspiracular bands margin pores between with spiracles, (3)particularly large on the surface; dorsal setae Alecanopsis Bombacoccus Williams & Watson (1990);Murphy (1997) Ben-Dov Ben-Dov Murphy (1997); Williams & Watson (1990) Green (1904b);Le Pelley (1968); Mamet (1950); (2013); Culik (1926);Barrera Ballou (2008);Ben-Dov Abrahao &Mamprim (1958);Almeida (1973); Watson (1990) Nakahara (1981);Shafee (1904b, 1909); Le Pelley (1968); Murphy (1997); Almeida (1973);Ben-Dov Ben-Dov Ben-Dov (1968); Murphy (1997) Ben-Dov Ben-Dov (2013) Adenuga &Adeboyeku (1987);Ben-Dov Present study Le Pelley (1968);Murphy (1997) Muzaffar & Ahmad (1977) LottoDe (1965);Le Pelley (1968);Murphy (1997); Pelley (1968);Murphy (1997),Shafee Ali (1971);Ben-Dov Cockerell, et al et al et al et al et al Kondo, Akermes . (2013); De Lotto. (2013);De (1968) . (2013);Hanford (1974) . (2013);Hanford (1974) et al . (2013); De Lotto. (2013);De (1963);Le Pelley . (2013);Nakahara (1981); &Miller . (2010); De Lotto. (2010);De (1956,1965); References et al Cockerell, Cribrolecanium Aztecalecanium . (2013); De Lotto. (2013);De (1956);Le et al et al . (1989); Williams & . (2013);Green et al Alecanium . (1989) Kondo Green, et al et al . .

GENÉTICA VEGETAL p. 39-51

Y BIODIVERSIDAD 47 A new species of Toumeyella Cockerell (Hemiptera: Coccoidea: Coccidae) on coffee roots,Coffea arabica L. (Rubiaceae), from Colombia and Venezuela enero - junio / 2013 GENÉTICA VEGETAL p. 39-51

Y BIODIVERSIDAD 48 A new species of Toumeyella Cockerell (Hemiptera: Coccoidea: Coccidae) on coffee roots,Coffea arabica L. (Rubiaceae), from Colombia and Venezuela enero - junio / 2013 of first-instar nymphs, Kondo and (2002) Williams on thehave multiple morphology evolved based times; 2009). It appears thatlimbs with softscales reduced synonym of It that noted be should Kondo & Williams 2002; Kondo 2010a; Octolecanium known to occur on the roots crown androot ofits host, coffeae (Kondo &Pellizzari,Regions 2011). found in other countries in the Nearctic and Neotropical that suggesting USA, many be should more species new New World with from being described most the species genusThe dataCO1 (T. unpublished data). Kondo &L.G. Cook, 28Sandpartial using on 18S, DNA based sequences a Bayesian phylogenetic analysis oftheCoccidae family unrelated This also is by supported hypothesis lineages. that the Myzolecaniinae ofseveral iscomposed suggested Adenuga AO, Adeboyeku 1987.Notes K. on distribution ofant- Abrahao J, Mamprim O. docafeeiro. da raiz 1958. Cochonilha O REFERENCES Bombacoccus Cyclolecanium the Toumeyella-group of iscomposed pores in Currently,quinquelocular the perivulvar region. and(4)presence of with setae; anal region hypopygial stigmatic setae small and inconspicuous or absent; (3) but retaining and (2)marginal reduced, their parts; theby following andantennae features: (1)legs greatly Steinweden defined the (1929)chiefly Toumeyella-group genera the Toumeyella-group,called in the heincluded which limbsformedwith a natural reduced group, he which Steinweden (1929)postulated that New World species Kondo 2011;Kondo2009; Kondo &Williams 2010a, 2004a, Torarchus Matile-Ferrero Ruyet, &Le Takahashi, Neolecanium Megasaissetia Kondo, Cryptostigma and its Application 8:239-243. Homoptera interaction crop on plants. selected Insect Science Sao PauloBiológico. 24:268-271. is the only species in isthe the species only genus et al et Toumeyella Halococcus Gullan &Stewart (Gullan &Stewart, 1996; Toumeyella ., 2005). Parrott, Toumeyella Ferris, Pseudophilippia , Cockerell, Pseudophilippia , Cyclolecanium Octolecanium Takahashi, Neolecanium has awide distribution in the by Williams &Kondo (2004b, Neolecanium , Megasaissetia Cockerell, Toumeyella Myzolecanium and Morrison, Kondo, and Houardia Toumeyella isconsidered a Pseudophilippia Cockerell and Paractenochiton , Neolecanium Foldilecanium et al et Richardiella Toumeyella Toumeyella Beccari, Beccari, Marchal, Akermes (Kondo ., 2005). , . ,

Musa sapientum, M. textilis guineensis hosts including Panama isknown from the system root ofvarious Kondo Ecuador Costa and from Rica, Colombia, 1914),and &Piza, Page 1900, Le and isanimportant ofvines in (Hempel, Brazil restrictedto be to the underground vines roots ofgrape Cryptostigma silveirai that on couple ofspecies thea roots, i.e., feed are there the system root oftheir hosts. Infrom the Myzolecaniinae, personal observation). Very other are few known coccids species recorded from coffee fromin recorded coffee species Cuba. Novoa for providing information on the reviewing the manuscript, andto Mrs. Nereida Mestre- Nationallian University, Canberra, Australia) for kindly USA)Alabama, andDr. Penny J. Austra Gullan (The 1. Many thanks to Dr. Nate Hardy (Auburn University, Fernanda Niño Díaz for providing the photos in Figure thanks andMaría to Special Clemencia Villegas studied. Niño Urtiaga andRadamés for the collecting material Jáuregui, Manuel G.,María Alexis Leal Fernanda Díaz coffeae (ICA) for providing the Venezuelan specimens of The author Ramosthanks Andrea Amalia Portilla ACKNOWLEDGEMENTS personnel. researchers workingas pestsquarantineas well on coffee for The presentinformation paper provide should useful officinale Zingiber (), closely related genus genus closely related lattertwoThe arecommon featuresantennae. in the the claw,the segmentation thefor poor of and(iii) except the segments complete of the of all fusion legs pores, (ii) pores that are about the the samesizeas spiracular the small sizeofthe perivulvar (i) its congeners by: all oftheirparts hosts. Morphologically, itdiffers from other all knownas are known species from the aerial Museum Bulletin 6:7-82. ofthe Oriental (Part 1971.Acatalogue SM. Ali Coccoidea V) 1968.Description andrecords ofsome knownSM. ofanew coccids Ali Aitken-Soux P. 1985.Quelques pestes généralement etmaladies (Insecta: Homoptera: Coccoidea) (with Homoptera: anindex). Coccoidea) (Insecta: Indian from Bihar, India. Oriental(Homoptera) Insects. New 1:29-43. Delhi Interaméricain deCoopération pour l’Agriculture, Haïti 53:1-5. dans lespépinièresrencontrées decafé. Feuille d’Extension, Institut . Many thanks Fabio to Clemencia Alonso Villegas, (Arecaceae), Theobroma (Zingiberaceae) (Kondo, 2010b). 2010b). (Kondo, (Zingiberaceae) Revista Corpoica -Ciencia y Tecnología Agropecuaria (Hempel), a species which appears which (Hempel), aspecies Akermes (Musaceae), sp. (Araceae), sp. (Sterculiaceae) and Cockerell (T. Kondo, sp. (Bromeliaceae), C. rhizophilum Peristeria elata Toumeyella Elaeis T. -

© 2013Corporación Colombiana deInvestigación Agropecuaria LottoG.1965.On (Homoptera) someDe Coccidae from chiefly LottoG.1963.New genusDe ofhard andanew species from scales Coccoidea: in Africacoffee of green scales south LottoG.1960.The De (Homoptera: of scales soft New 1958. G. Lotto De LottoG.1957.On someDe Ethiopian ofthe species genus Coccus Coccoidea) Trudy (Hemipt.: LottoG.1956.TheidentityDe East ofsomeAfrican of species coccids Russian). new Three (In 1955. G. Vietnam]. Lotto De of fauna Coccinea) KonstantinovaDanzig EM, (Homoptera, 1990.[On coccid GM. PJ, Gullan ALBG, Peronti JA, Ventura DS, Martins MP, Culik Coccidos Chazeau J. de 1981.Données sur la biologie 1986. G. Guerra Pérez A, Hernández Carnero insects on SO. citrusBoboye 1971.Scale andtheir distribution in Ben-Dov Y. 1993.Asystematic ofthe insects softscale of catalogue Ben-Dov Y. 1979.Ataxonomic studygenus ofthe soft-scale Ben-Dov Y.genus of scales the 1970.The wax Barrera JF.pests and 2008.Coffee their management. Pp. 961-998, deCuba cóccidos ysus 1926.Los plantas CH. Ballou hospederas. ofthe genusAvasthi 1991.Revision SA. Shafee RK, especies das 1- Revisao de Angola. 1973. Coccoidea de DM. Almeida the Sahara (Homoptera, Coccidae). Bulletin of Entomological Entomological of 51:389-403. Research Bulletin Coccidae). (Homoptera, Sahara the (B)27:165-172. ofLondon Society ofthe Entomological Royal from Proceedings Coccidae) Africa. ofSouthern AfricaSociety 20:295-314. Journal Coccidae). of the Entomological Coccoidea: (Homoptera: 47:239-249. Research Entomological Saissetia of Bulletin Africa. East 46:267-273. Research in coffee attacking 209: 38-52. Instituta Leningrad Zoologicheskogo Akademiya Nauk SSSR. Santo, Espírito of BiotaBrazil. Neotropica 7(3):1-5. Coccoidea) (Hemiptera: Monophlebidae Kondo T. Pseudococcidae, and Ortheziidae, 2007.Coccidae, 301-311. (Hom.: en Coccidae) Nouvelle-Calédonie. Entomophaga 26: Serie: quadrivittata Agrarias, Investigaciones de Nacional Protección Vegetal. 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(Homoptera, Coccidae). Bulletin of Entomological Entomological of Bulletin Coccidae). (Homoptera, (Col.: ), predateur Coccinellidae), (Col.: viridis deCoccus Ceroplastes Coccus Coelophora Coelophora Linn., Linn., Kilifia Gray González-Hernández H, Atkinson TH. 1984. Cocoideos Cocoideos 1984. Granara Pirovani MC, deWillink VD, Ferreira es 2010.Las PS. TH. Atkinson H, González-Hernández Scales Scale RJ. Soft Gill 1988.The 1.The Part Insects of California: Fonseca 1957.Três da. JP doBrasil, decoccídeos novas espécies Fernández “escamas 1957.Las A. palomillas” delcafeto en Venezuela andmealybugs softscales andnew old LottoG.1969.On afew De contribution Lotto G.1968.Second De to ofthe the knowledge LottoG.1967.AcontributionDe to ofthe the African knowledge Hamon 1984.Thesoft ofscale insects Williams ML. Florida AB, of species Hall WJ. Bulletin,pests 1924.The ofcitrusinsect trees in Egypt. and genus new A 1996. AC. Stewart PJ, Gullan 1937.An annotatedGreen EE. with listofthe ofCeylon, Coccidae from on some Coccidae Zanzibar, 1916.Report collected EE. Green IV. ofCeylon. 1909.TheCoccidae Part Green EE. Dulau &Co., Notes by the 1907.XII. PercyGreen on EE. the collected Coccidae 1904b. On some JavaneseGreen EE. with descriptions Coccidae: Dulau &Co.,III. Pt. ofCeylon. TheCoccidae 1904a. Green EE. 1903.Remarks onGreen Indian EE. With insects scale (Coccidae). Indian in Ceylon. collected ofCoccidae 1896.Catalogue Green EE. 1889.Descriptions of Green species EE. new oftwo 39(3):391-399. Entomology Neotropical de dos especies. y Redescripción Coccidae) pecies de la pecies de frutales árboles a centralregión deMéxico. 57:207-225. Agrociencia asociados Coccoidea) (Homoptera: Food 132p. &Agriculture, Sacramento, USA. CA, California Coccidae). Department of (Homoptera: Coccoidea: São PauloBiológico, 24:123-135. sobre cafeeiro (Homoptera Arquivos -Coccidae). doInstituto (Venezuela) 7(1):7-17. y avancesen su control logrados químico. Tropical Agronomía of Southern Africa 32:413-422. Journal(Homoptera: Coccoidea). Society ofthe Entomological ofSouthern AfricaSociety 31:83-86. (Homoptera).African Coccoidea Journal ofthe Entomological of Southern Africa 29:109-120. (Homoptera).Coccoidea Journal Society ofthe Entomological FL, USA. 194p. USA. FL, & Consumer Division ofPlant Industries, Service, Gainesville, Neighboringand LandAreas. Florida Department ofAgriculture In: Coccidae). of Florida (Homoptera: Coccoidea: 45: 1-29. Technical ofAgriculture,Ministry Egypt, andScientificService Museum 39:307-14. from Myzolecaniinae) (Hemiptera: Coccidae: ant-associated coccid 20:277-341. andGeology B.Zoology Section emendations andadditions to date. Journal Ceylon ofScience Dr.by W.M. Aders. Bulletin 6:375-376. 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GENÉTICA VEGETAL p. 39-51

Y BIODIVERSIDAD 49 A new species of Toumeyella Cockerell (Hemiptera: Coccoidea: Coccidae) on coffee roots,Coffea arabica L. (Rubiaceae), from Colombia and Venezuela enero - junio / 2013 GENÉTICA VEGETAL p. 39-51

Y BIODIVERSIDAD 50 A new species of Toumeyella Cockerell (Hemiptera: Coccoidea: Coccidae) on coffee roots,Coffea arabica L. (Rubiaceae), from Colombia and Venezuela enero - junio / 2013 od T Wlim M, uln J 20. eecito of Redescription 2005. PJ. Gullan ML, Williams T, Kondo T,Kondo Williams DJ. designation 2008.Neotype andredescription of species new a of Description 2011. G. Pellizzari T, Kondo Kondo T, J, Reyes Monsalve Muñoz J, R, Mesa 2011. López NC. JA, Kirkaldy G.1902.Hemiptera. Fauna Hawaiiensis 3:93-174. Ihering (Phytophthires) H von.vegetaes 1897. Os piolhos do Brazil. anidentification Coccidae: CJ.family insect 1994.Thescale Hodgson CJ.Hodgson 1969b. Notes (Homoptera: Coccidae on Rhodesian od T 21a Dsrpin f nw ocd (Hemiptera, coccid new a of Description 2010a. T. Kondo T.Kondo genus, African 2006. A new soft scale Kondo T, 2002.The Williams (Hemiptera: ML. Myzolecaniinae Kondo T, 2009.Redescriptions Williams ML. of T,Kondo Redescription ofthe myrmecophilous 2004a. Williams ML. Kondo T, 2004b. Redescription Williams ofthe ML. Mexican (Homoptera: Coccidae Rhodesian on Notes 1969a. CJ. Hodgson CJ.Hodgson 1967.Notes (Homoptera: Coccidae on Rhodesian Hempel 1921.Três A. Archivos da Escola Superior novos coccídeos. Hempel 1900.Descriptions A. ofthree ofCoccidae species new Hanfordgenus 1974.The African L. insect scale Octolecanium perconvexum Coccidae) on avocadoCoccidae) ( nov. Journal ofInsect 6(1):1-16. Science includingspecies, the citrus pest for erected two nov. Coccidae), gen. (Hemiptera: Coccoidea: Ser. Bachicoltura, edi Agraria 33(3):125-128. II, OldWorldCoccidae): vs.New World. Zoologia Bolletino di America. South Colombia, and USA International Journal ofInsect 2:11-27. Science southeastern the from with(Hemiptera: Coccidae), descriptions species new oftwo Ckll. &Parrott andtheir transfer to leucaenae 215. Ann.Hemiptera: Coccidae). Naturhist. Mus. Wien. B105:211- insect soft scale 7(1):5-9. en Ciencias Especializada Revista Químico-Biológicas TIP Coccidae). combinationand new (Hemiptera: Coccoidea: insect: soft scale from Species New a 8(1):5-11. en CienciasEspecializada Químico-Biológicas of Description Revista TIP Coccidae). Guatemala (Hemiptera: Coccoidea: with Combination, JournalCoccidae). ofInsect 8(56):1-6. Science of 55(2):229-233. Entomologia with ataxonomic key to Mexican Brasileira Revista de species. Toumeyella 20 p. y elValle Produmedios, Bogotá. Corpoica. delCauca, Colombia. Insectos escama yácaros comunes en elEje delaguacate Cafetero doMuseuRevista Paulista. São Paulo 2:385-420. 639 p. International,manual CAB to genera. Wallingford, Oxon, UK. Part Arnoldia 4:1-42. (Rhodesia) Coccoidea): III. Arnoldia 4:1-43. (Rhodesia) genera The PartCoccoidea): II: and a new genus and anew PartgeneraCoccoidea): 1: The de Agricultura eMedicina Veterinária. RiodeJaneiro 5:143-146. from Canadian Brazil. Entomologist 32:3-7. 126:1-40. London Transactions(Coccidae). of Society ofthe Entomological Royal Toumeyella liriodendri Ckll., Cockerell from (Hemiptera: Coccidae) Mexico; Aztecalecanium colimae Toumeyella sallei Mashona Toumeyella cerifera Persea americana . Arnoldia 3:1-22. (Rhodesia) (Gmelin) (Hemiptera: Coccoidea: (Gmelin) (Hemiptera: Coccoidea: (Cockerell), New(Cockerell), Genus and New P. andersoni (Signoret), comb.n. (Insecta: (Insecta: comb.n. (Signoret), Coccus Ceroplastes Ferris and Neotoumeyella (Cockerell), new genus new (Cockerell), , Mill.) from Colombia, from Mill.) Colombia, Parasaissetia Pseudocribrolecanium (Newstead) comb. Udinia and T. sonorensis Neolecanium Gascardia De Lotto De e. nov. gen. , Saissetia .

ae J. 99 Nts n h Ccoda f aaacr IV. Madagascar Mestre Novoa N, Hamon Evans A, G,Kondo T, of Oliver P, Marrero Coccoidea the on Notes 1959. JR. Mamet andother 1997.First IL. reportLit ofthe Lecanodiaspididae family Piza MT. HS, Lepage do 1941.Redescrição Pelley 1968.PestsLe RH. Longmans, 588 London. p. ofcoffee. Laing F. 1929.Descriptions ofnew, of andsome notes onspecies old Kondo T. 2011. Transfer of the myrmecophilous softinsect scale myrmecophilous, the of revision Taxonomic 2010b. T. Kondo Shafee SA, Yousuf SA, Shafee Khan M, MY. 1989.Host plants anddistribution Ramakrishna Ayyar TV. 1930.Acontribution to our knowledge Ramakrishna Ayyar TV. 1919. A contribution to our of knowledge 1920. Observations on - VI. scale-insects (Coccidae) Newstead R. 1917.Observations on -V. scale-insects (Coccidae) Newstead R. andAleurodidae, ofCoccidae 1911.On acollection R. Newstead species Coccoidea Nakahara the ofthe S.1981.List Hawaiian (Homoptera: Coccoidea of list A 1981. CE. Miller S, Nakahara MuzaffarR. 1977. A note on N, Ahmad Y. & Ben-Dov In: 367-380. Coffee. 3.3.14 1997. ST. Murphy Murphy ST. 1991.Insect naturalgreen scales enemies ofcoffee meliponiphilous and rhizophilous soft scale genus andrhizophilous softscale meliponiphilous 42:37-45. Keys South America. Zoo Coccoidea) presentesCoccoidea) en la Cordillera Pinar deGuaniguanico, (Hemiptera: : cocoideos 2011.Los Alonso A. A, 369-479. Mémoires del’Institut Scientifique (Ser. de Madagascar E)11: Hemiptera). Philippine Entomologist 11:87-95. records andnotes onnew Philippine insects scale (Coccoidea, controle. Arquivos doInstituto São Paulo Biológico. 12:21-26. da(Hempel) videira séria (Homoptera-Coccoidea), eseu praga AnnalsCoccidae. ofNatural andMagazine 4:465-501. History InsectaColombia. Mundi 0167:1-10. with(Hemiptera: Coccidae), description from species ofanew Neolecanium amazonensis 2709:1-72. Zootaxa Coccidae). Ferris (Hemiptera: Coccoidea: of Systematic 6:47-55. Entomology in India. pests Indian (Homoptera: Coccoidea) of coccid Journal 1-73. the Imperial Institute of Agricultural India Pusa, Research, 197: Bulletinof South Indian andMealybugs). of (Scales Coccidae Institute, India Pusa, 87:1-50. South Indian Bulletin Coccidae. of the Agricultural Research Bulletin 10:175-207. Research ofEntomological Bulletin 8:125-134. Research ofEntomological 155-174. Museum ausMitteilungen dem Zoologischen in Berlin 5: seum. chiefly African,Mu in of Zoological collection Berlinthe the 23:387-424. Society Sternorhyncha). ofthe Hawaiian Proceedings Entomological ofWashingtonSociety 83:28-39. (Homoptera) ofthe ofPuerto Entomological Rico. Proceedings Entomophaga 22:45-46. in PakistanCoccidae) andthe ofits natural breeding enemies. York. 442p. Enemies and Control [Vol. 7B]. Elsevier, Amsterdam & New Scale (Editors), Soft Insects -Their Biology, CJ Natural Hodgson Entomophaga 36:519-529. control of in Kenya Coccidae) (Hemiptera: and their potential for biological 0183: 1-25. yladel Río,Cuba, relación con sus hospedantes. Insecta Mundi Coccus celatus Revista Corpoica -Ciencia y Tecnología Agropecuaria and Foldi to Coccus viridis Foldilecanium Saissetia privigna Saissetia in Papua New Guinea. Neolecanium silveirai Cryptostigma e. nov. gen. (Hem.: -

© 2013Corporación Colombiana deInvestigación Agropecuaria Williams DJ.distribution 1982. The synonymy and of Williams DJ, Watson GW. Scale 1990.The Insects of the Tropical Vayssière P. animaux 1955.Les du parasites caféier. (In French). p. Steinweden for 1929.Bases the JB. generic classification of the Silva d’Araujo DM. Goncalves Galvao GM, CR, Goncalves GA, BJ,ShefferDescriptions, 1990. ML. Williams distribution, and 107-109. in Papua New Bulletin 72: Research Guinea. ofEntomological andits Lotto(Hemiptera: importance Coccidae) De on coffee International, CAB other families. Wallingford, 267p. UK. South (Coccidae) and Pacificscales 3.Thesoft Part Region. Monde, Paris. 382p. Caféiers (Editor), Les etlesCafé257-267. In: dans Coste le R. of America 22:197-245. Annals Coccidae. family Society ofthe Entomological coccoid V. 1,622p. hosts andnatural enemies. Ministerio da Cultura, RiodeJaneiro. 1968. Fourth catalog ofinsects that live in Part Brazil. Insects, II. Entomological the of ofWashingtonSociety 92(1):44-57. Proceedings Coccidae). (Homoptera: host-plant first records ofeight instarsgenus in the Coccus celatus Toumeyella

Wolcott 1941.Supplement GN. to 2010. Annotated ML. listof the insects scale of Guatemala.Williams 1993. Williams ML. Kondo T.Williams ML, 2008.Status andcurrent composition of nymphs first-instar of Characteristics 2001. T. Kondo ML, Williams Williams on DJ. coffee insects (Homoptera: Coccoidea) 1986.Scale Agriculture ofthe University ofPuerto 25:33-158. Rico HellenicaEntomologia 19:144-152. Entomologist 76(4):566-572. insectfrom the Floridascale keys Florida (Homoptera: Coccidae). Portugal. SeptemberOeiras, 24-27,2007.Pp. 29-32. Insect Branco Franco M, Studies. Editors: C. &Hodgson JC of the XIthProceedings International Symposium on Scale genusthe softinsect scale surprising Coccidae): (Hemiptera: Bachicoltura (Milano) 33(3):35-42. insects scale e di Agraria indicators of relationships. Bollettino Zoologia di soft the in Forestry andFisheries 34:1-7. Papuain New Papua Guinea. New Guinea Journal ofAgriculture, Toumeyella lignumvitae Toumeyella Insectae borinquenses (Hemiptera: Coccoidea). (Hemiptera: Coccoidea). a e seis of species new a , . Journal of

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Y BIODIVERSIDAD 51 A new species of Toumeyella Cockerell (Hemiptera: Coccoidea: Coccidae) on coffee roots,Coffea arabica L. (Rubiaceae), from Colombia and Venezuela enero - junio / 2013