Mealybugs of the Genera Planococcus and Crisicoccus (Sternorrhyncha: Pseudococcidae) of Russia and Adjacent Countries

ZOOSYSTEMATICA ROSSICA, 19(1): 39–49 15 JULY 2010

Mealybugs of the genera Planococcus and Crisicoccus (Sternorrhyncha: Pseudococcidae) of Russia and adjacent countries Мучнистые червецы родов Planococcus и Crisicoccus (Sternorrhyncha: Pseudococcidae) фауны России и сопредельных стран

Е.М. DANZIG & I.A. GAVRILOV

Е.М. ДАНЦИГ, И.А. ГАВРИЛОВ

Е.М. Danzig, Zoological Institute, Russian Academy of Sciences, 1 Universitetskaya Emb., St. Petersburg, 199034, Russia. E-mail: [email protected] I.A. Gavrilov, Zoological Institute, Russian Academy of Sciences, 1 Universitetskaya Emb., St. Petersburg, 199034, Russia. E-mail: [email protected]

A key for four species of the genus Planococcus Ferris, 1950 is given; three species inhabit the territory of Russia and adjacent countries and the fourth one, P. citri (Risso, 1813), was errone- ously recorded from the territory of the former USSR, because of the confusion of this species with P. ficus (Signoret, 1875) during identification. All discussed species are morphologically described and illustrated. Planococcus taigae Danzig, 1980 and P. juniperus Tang, 1988 are placed in synonymy under Planococcus vovae Nasonov, 1908. The genus Crisicoccus Ferris, 1950, which is morphologically similar to Planococcus, is also discussed and reported for the territory of Russia for the first time with the species C. pini (Kuwana, 1902). В статье приводится определительная таблица для 4 видов рода Planococcus Ferris, 1950; 3 из них обитают на рассматриваемой территории, четвертый, P. citri (Risso, 1813), ранее указывался ошибочно в связи со сложностью его дифференциации от P. ficus (Signo- ret, 1875). Для всех обсуждаемых видов даны рисунки и описания. Planococcus taigae Danzig, 1980 и P. juniperus Tang, 1988 помещаются в синонимы Planococcus vovae Na- sonov, 1908. Рассмотрен также ближайший к Planococcus род Crisicoccus Ferris, 1950, представленный в России единственным видом C. pini, который впервые указывается для рассматриваемой фауны. Key words: scale insects, Pseudococcidae, Planococcus, morphology, taxonomy, new synonyms Ключевые слова: кокциды, Pseudococcidae, Planococcus, морфология, таксономия, новые синонимы

TAXONOMIC PART Allococcus Ezzat & McConnell, 1956: 13 (type species Pseudococcus inamabilis Hambleton, Order STERNORRHYNCHA 1935, by original designation). Family PSEUDOCOCCIDAE Allococcus: Cox & Ben-Dov, 1986 (= Planococcus). Planococcus Ferris, 1950 Type species: Pseudococcus citri Risso, 1813, by original designation. Planococcus Ferris, 1950: 164. Diagnosis of the genus. Adult female. Planococcus: Ezzat & McConnell, 1956: 60; Body oval. Anal lobes with characteristic McKenzie, 1967: 280; Danzig, 1980: 168; anal lobe bars. Antennae 8-segmented. Legs Cox, 1989: 1–78. well developed; coxae, tibiae and sometimes

© 2010 Zoological Institute, Russian Academy of Scienсes1 40 Е.М. DANZIG & I.A. GAVRILOV. PLANOCOCCUS AND CRISICOCCUS OF RUSSIA AND ADJACENT COUNTRIES femora of hind legs with translucent pores. on hind femora. Multilocular pores present Circulus usually present. Spiracles well de- on last 5 abdominal segments; sometimes veloped. Multilocular pores numerous on single pores present also on thorax. Single ventral parts of abdominal segments; some- dorsal tubular ducts present near each times single pores present on ventral parts cerarii; ventral tubular ducts form trans- of thorax and head and also on dorsum. Tu- verse rows in the middle part of abdomen bular ducts present on both body sides; usu- and groups along abdominal margin; sin- ally dorsal ducts larger and sometimes with gle ducts present in middle part of thorax; collars. Cerarii usually totaling 18 pairs; sometimes 1–5 ducts present on head and sometimes thoracic cerarii absent, but С3 near first thoracic cerarii. Eighteen pairs of always well developed. All cerarii with two cerarii present; setae of head cerarii with conical setae (sometimes cerarii on head flagellate apex. with 3 conical setae); additional flagellate Remarks. The two sympatric species, P. setae present in С18 only. С18 are situated on ficus and P. citri are very similar to each oth- sclerotized plates. er morphologically and ecologically. Some The genus comprises about 40 species, authors (Borchsenius, 1949; Ferris, 1950; mainly tropical and subtropical in distribu- McKenzie, 1967; Ter-Grigorian, 1973; De tion. Only P. vovae (Nasonov, 1908) and P. Lotto, 1975; Terznikova, 1975; Khadzibeili, matesovae Danzig, 1986 inhabit the boreal 1983) regarded these as a single species. In zone. Many species of the genus are impor- recent coccidological papers P. ficus and tant pests of agricultural and ornamental P. citri are considered as separated species. plants; in the fauna of Russia and neighbor- During the study of material from the terri- ing countries P. ficus (Signoret, 1875) is tory of former USSR we verified that only P. the only economically important species of ficus inhabit the discussed territory. The re- the genus. cord of Danzig (1977) of P. citri in Abkhazia Most species of the genus Planococ- (Sukhum) on Vitis sp., Ficus carica and Pu- cus are very similar morphologically and, nica granatum was based on the presence of at the same time, demonstrate significant two groups of tubular ducts (with 2–8 ducts variability. in each group) on head of some studied individuals. However, the other individuals Planococcus ficus (Signoret, 1875) from the same populations did not have such (Fig. 1) groups. So, this character varies between Dactylopius ficus Signoret, 1875: 315 (France). specimens. Cerarian setae of all analyzed fe- Planococcus ficus: Ezzat, McConnell, 1956: 79; Dan- males from Abkhazia are thin and with flag- zig, 1977: 99; Cox, Ben-Dov, 1986: 483; Cox, ellate setae. Also, three females with similar 1989: 25; Williams, Moghaddam, 2000: 34. morphological characters were found by us Coccus vitis (non Linnaeus) Nedzelskii, 1869: 19 in material collected by E.F. Kozarzevskaya (misidentification). in greenhouses of the Main Botanical Gar- Pseudococcus citri auct. (non Risso, 1813): den on Euphorbia and Coleus. Borchsenius, 1949: 131; Ter-Grigorian, 1973: Differential characters between P. ficus 82; De Lotto, 1975: 125; Tereznikova, 1975: and P. citri are noted in the Key (see below). 226; Khadzibeili, 1983: 67. Both of these species probably originated Material. Thirty slides from greenhouses of from Mediterranean region and spread to the Main Botanical Garden, Moscow. More than 100 slides from Spain, France, Ukraine (Crimea), other regions through the time with the cul- Abkhazia, Georgia, Armenia, Azerbaijan, Turkey, tivation of grapes and Citrus. Now both spe- Turkmenistan, Uzbekistan, Egypt, Israel, Iraq, cies are distributed everywhere in tropical Afghanistan, India, Vietnam, USA. and subtropical regions and in greenhouses. Description. Adult female. Translucent It seems that both species have the same hy- pores present on hind coxae and sometimes menopteran parasitoids, but, for example, in

Fig. 1. Planococcus ficus, Uzbekistan, from Morus.

Israel some P. ficus parasitoids were never Pseudococcus (Dactylopius) vovae Nasonov, 1908: noted on P. citri (Noyes & Hayat, 1994). Se- 484 (Poland). rological methods also do not show any dif- Pseudococcus vovae: Borchsenius, 1949: 134. ferences; and moreover, crossings between Pseudococcus junipericola Borchsenius, 1949: P. ficus and P. citri are possible (Tremblay et 116 (Tajikistan). Pseudococcus junipericola: Danzig, 1986: 18 (syn- al., 1980; Tranfaglia & Tremblay, 1982). onymy, lectotype designation). Experiments in which P. ficus is grown Pseudococcus inamabilis Hambleton, 1935: 112 under different temperatures showed that (Brasil). the translucent pores of the hind femora (a Allococcus inamabilis (Hambleton, 1935): Ez- taxonomically important character) were zat & McConell, 1956: 15; Danzig, 1986: 18 present in high temperature and absent in (synonymy); Roberti, Tranfaglia, 1981: 305. low temperature (Cox, 1989). Planococcus taigae Danzig, 1980: 168 (P. vovae, Distribution. Russia (greenhouses of the partly); 1986: 19 (Russia), new synonym. Main Botanical Garden, Moscow); Ukraine Planococcus taigae: Cox, 1989: 68. Planococcus juniperus Tang in Tang & Li, 1988: (the Crimea), Transcaucasia, Middle Asia, 42 (China, Inner Mongolia), new synonym. Mediterranean region, the Near East; intro- Planococcus juniperus: Tang, 1992: 352. duced in India, Pakistan, Vietnam, Argen- tine, Brazil, USA (California), South Africa. Material. Russia (Leningrad Prov., Karelia, Mode of life. Polyphagous; inhabit over- North Caucasus, Siberia, South of Primorsk Ter- ground parts of numerous wild, agricultural ritory, South Sakhalin, Kuril Islands (Shikotan); Ukraine (Crimea and Odessa Prov.), Armenia, and ornamental plants; most common on mountains of Middle Asia (including the type Vitis, Ficus, Punica granatum, Platanus, series of P. junipericola), Spain, France, Czech and Morus. In the Crimea and Caucasus the Republic, Poland; in general 64 slides. species has three generations per year (Um- Description. Adult female. Circulus pres- nov, 1940; Khadzibeili, 1985). ent (sometimes absent in insects from the mountains of Middle Asia). Multilocular Planococcus citri (Risso, 1813) pores located on last five abdominal seg- (Fig. 2) ments. Number of dorsal tubular ducts Dorthesia citri Risso, 1813: 416. varies geographically (see Remarks); some- Synonymy: see Ben-Dov, 1994: 355. times some tubular ducts appearing to be of the oral rim type. Ventral tubular ducts nu- Material. Series of females from Iran, India, merous on all segments; form groups along China, Vietnam. body margin; marginal ducts a little bit Remarks. The description of Borchse- larger than median ones. Cerarii 18 pairs in nius (1949) includes mixed characters of P. number; sometimes 1 or 2 pairs of thoracic citi and P. ficus. Detailed descriptions of P. cerarii absent. Cerarian setae (excluding citri, its differences from similar species and С ) in females from Spain, France, Georgia experimental data on interspecific variabil- 18 ity are given in the papers by Cox (1981, and Crimea, collected from Cupressus and 1983, 1989). Juniperus are thin, long, flagellate or even Distribution. Polyphagous. Widely dis- hair-like. In the eastern part of their distri- tributed in the tropics and subtropics and bution, insects have conical, comparatively in greenhouses in the entire world; a serious short cerarian setae, but the cerarian setae pest of Citrus. on the head can be thinner and longer than the thoracic and abdominal ones. Remarks. Planococcus taigae differs from Planococcus vovae (Nasonov, 1908) P. vovae in the large number of dorsal tu- (Fig. 3) bular ducts and in having thicker cerarian Coccus gossipifera Rondani, 1874: 43, nomen setae. However, the first character varies oblitum. geographically: insects from the Mediter-

Fig. 2. Planococcus citri, quarantine material of Main Botanical Garden, Moscow.

Fig. 3. Planococcus vovae, holotype of P. taigae, after Danzig (1980). ranean region have a small number of ducts the Eastern part of the area; in general, in- in the last abdominal segments; insects sect populations from the Russian Far East from central Europe have much more ducts have thicker setae and numerous ventral tu- which form transverse rows on the entire bular ducts along the body margin. body (Cox, 1989; Williams & Moghaddam, Borchsenius (1949) separated P. juniper- 2000; Marotta, 1990); insects from North- icola from P. vovae by the presence of a large West Russia, Siberia and Far East (types of circulus in the former one. We have revealed P. taigae) have the greatest number of these that this character varies between individu- ducts. As for the thickness of cerarian setae, als; insects without a circulus are very rare this character varies amongst specimens in and were collected in the mountains of

Middle Asia only. Females from Eastern Remarks. The species is similar to P. vo- Turkey have even up to 2 circuli (B. Kay- vae, but differs in having more numerous dan’s material). The number of multilocular tubular ducts that are of two sizes and in the pores along the ventral abdominal margin presence of multilocular pores on dorsum. is also a variable character; these pores are Distribution. Kazakhstan: Kalbinskii and usually absent or rare, but specimens from Zailiiskii mountain ranges. Iran have many of these pores (Williams & Mode of life. The species was collected Moghaddam, 2000). from Juniperus sabina in June, on steppe The figure of P. juniperus drawn by Tang rocks. & Li (1988), shows no differences with P. vovae (=P. taigae) collected from the same Crisicoccus Ferris, 1950 species of Juniperus. Mode of life. Oligophagous, on Cupres- Crisicoccus Ferris, 1950: 45. saceae; inhabit branches and needles; some- Crisicoccus: Ezzat, McConnell, 1956: 22; Wil- times causing damage. Oviposition in July- liams, 1985: 94, 2004: 129; Williams & Wat- August. son, 1988: 41; Tang, 1992: 347. Distribution. Planococcus vovae is a Type species Dactylopius pini Kuwana, widely distributed Palaearctic species, pres- 1902, by original designation and monotypy. ent in Russia, from Western to Eastern bor- Remarks. The genus Crisicoccus was de- ders, Transcaucasia, mountains of Middle scribed by Ferris (1950) in the same mono- Asia, Europe, the Middle East, Turkey, Iran; graph with Planococcus during a revision of introduced in Brazil and Argentina. the American fauna and included a single species C. pini, introduced from Asia. Ferris Planococcus matesovae Danzig, 1986 (1950) also noted that C. pini could be prob- (Fig. 4) ably transferred to the genus Planococcus. Planococcus matesovae Danzig, 1986: 21 (Ka- Other authors also noted the close similar- zakhstan: Kalbinskii mountain ridge). ity of both discussed genera, in particular, Crisicoccus matesovae: Cox, 1989: 70. the presence of anal lobe bars. The main dif- Material. Types and series of females from ferences between Planococcus and Crisicoc- Zailiiskii Alatau (Kazakhstan). cus is a smaller number of cerarii, in particu- Description. Adult female. Multilocular lar the absence of C3 in Crisicoccus, which is pores on venter present on last five poste- well developed in Planococcus (Williams, rior abdominal segments; single pores pres- 2004). Currently, Crisicoccus comprises ent on IV–VII abdominal segments. Dorsal about 30 species, mainly from Asia and the tubular ducts of two sizes, forming trans- Australasian region, but their relationship verse bands on all segments, excluding last with the type species is not obvious. abdominal segment. Number of larger ducts decreases towards anterior part of body; Crisicoccus pini (Kuwana, 1902) some ducts appearing to be of the oral rim (Fig. 5) type. Ventral tubular ducts numerous on all segments; forming groups along body Dactylopius pini Kuwana, 1902: 54 (Japan); margin; marginal ducts slightly larger than Crisicoccus pini: Ferris, 1950: 46; Ezzat & McCo- median ones. Number of cerarii varies geo- nell, 1956: 29; McKenzie, 1967: 127. graphically: insects collected from the Kal- Material. One series of females from Khasan binskii range have 5–10 pairs (thoracic and region of Primorsk Terr. (Russia) and one else anterior abdominal cerarii absent); insects series from China (Tsindao). from Zailiiskii Alatau have 18 pairs of cer- Description. Adult female. Circulus ab- arii. Cerarian setae thin, flagellate, exclud- sent. Multilocular pores located in middle ing С18 which has more conical setae. areas of the last five ventral abdominal

Fig. 4. Planococcus matesovae, after Danzig (1986).

Fig. 5. Crisicoccus pini, Primorsk Territory.

© 2010 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 19(1): 39–49 48 Е.М. DANZIG & I.A. GAVRILOV. PLANOCOCCUS AND CRISICOCCUS OF RUSSIA AND ADJACENT COUNTRIES segments. Tubular ducts present on venter in number. Widely distributed in the Palae- of abdomen only; these ducts of two sizes: arctic region ...... P. vovae larger ones form groups along body margin, 6(5). Dorsal tubular ducts of two sizes. Multiloc- excluding 8-th abdominal segment, where ular pores present on dorsum. Sometimes first abdominal and thoracic cerarii absent. Known they are located in the middle area of the from Kazachstan only ...... P. matesovae segment; smaller ones located in middle areas of posterior ventral abdominal segments. ACKNOWLEDGEMENTS Cerarii 6–7 pairs in number; cerarian setae conical, rather thin. С18 with two additional The authors thank T. Kondo for valuable setae and a group of trilocular pores. Body remarks and linguistic revision. The work of covered by long thickened hairs. I.A.G. was supported by the Russian Foundation Remarks. The notable character of the for Basic Researches (projects No. 08-04-00787 species is the absence of a circulus. and No. 09-04-91229-Ста) and by the grant Distribution. Russia: Primorsk Terr. Ko- of the President of Russian Federation (МК- 6075.2010.4). rea, China (including Tibet), Japan; introduced to California (USA). Mode of life. The species inhabit the nee- REFERENCES dles of different Pinus species. In Primorsk Ben-Dov, Y. 1994. A systematic catalogue of the territory young females were collected from mealybugs of the world (Insecta: Homoptera: Pinus funebris in mid June. In California, Coccoidea: Pseudococcidae and Putoidae) it is known to cause damage to ornamental with data on geographical distribution, host pine-trees. plants, biology and economic importance. An- dover, UK. 686 p. Borchsenius, N.S. 1949. Homoptera. Suborder Key to species of the genus Planococcus Coccoidea. Family Pseudococcidae. Vol. 7. 1(4). Dorsal tubular ducts, if present, located Fauna SSSR. New Series. No. 38. Moscow– near cerarii only; sometimes 1 or 2 ducts Leningrad. 382 p. (In Russian). Cox, J.M. 1981. Identification of Planococcus present in middle part of abdomen. Most citri (Homoptera: Pseudococcidae) and the cerarii with conical setae, but cerarii on description of a new species. Systematic En- head and thorax sometimes thin, with flag- tomology, 6: 47–53. ellate apex. On deciduous trees and shrubs. Cox, J.M. 1983. An experimental study of 2(3). Tubular ducts on ventral part of head and morphological variation in mealybugs (Ho- near first thoracic cerarii absent; if pres- moptera: Coccoidea: Pseudococcidae). Sys- ent, they are located on head and thorax, in tematic Entomology, 8: 361–382. groups of 1–5 ducts in each group. Cerarii on Cox, J.M. 1989. The mealybug genus Planococ- head and thorax with flagellate apex. Hind cus (Homoptera: Pseudococcidae). Bulletin femora usually with translucent pores ...... of the British Museum (Natural History). En- ...... P. ficus tomology, 58(1): 1–78. 3(2). Tubular ducts on ventral part of head and Cox, J.M. & Ben-Dov, Y. 1986. Planococcine thorax present; up to 30 ducts in a single mealybugs of economic importance from group. All cerarii with conical setae. Some- the Mediterranean Basin and their distinc- times setae in head pairs of cerarii thinner. tion from a new African genus (Hemiptera: Translucent pores absent on hind . . . .P. citri Pseudococcidae). Bulletin of Entomological 4(1). Dorsal tubular ducts numerous (13–200 in Research, 76: 481–489. number); located along body margin and in Danzig, E.M. 1977. On the nomenclature and middle areas of body. All cerarii with thin set- distribution of some injurious scale insects ae; although some cerarian setae may be thick (Homoptera, Coccoidea). (In Russian). En- in individuals from the Eastern part of the tomologicheskoye Obozreniye, 56: 99–102. species range. On Juniperus and Cupressus. (In Russian). 5(6). Dorsal tubular ducts of one size. Multiloc- Danzig, E.M. 1980. Koktsidy Dalnego Vostoka ular pores absent on dorsum. Cerarii 18 pairs SSSR (Homoptera, Coccinea) s filogeniey

koktsid mirovoy fauny [Scale insects of the départment des alpes maritimes. Annales du Far East USSR (Homoptera, Coccinea) with Muséum National d’Histoire Naturelle, 20: phylogenetic analysis of scale insects fauna of 401–431. the world]. Leningrad. 367 p. (In Russian). Roberti, D. & Tranfaglia, A. 1981. Allococcus Danzig, E.M. 1986. New data on the systemat- inamabilis (Hambleton) on Cupressus mac- ics of Coccinea (Homoptera) of the USSR rocarpa, new to Italy. Memorie della Societа fauna. Vestnik Zoologii, 4: 18–22. (In Russian Entomologica Italiana, 60: 305–309. with English summary). Rondani, C. 1874. Degli insetti nocivi e dei loro De Lotto, G. 1975. Notes on the vine mealybug parassiti. Bollettino della Societа Entomologi- (Homoptera: Coccoidea: Pseudococcidae). ca Italiana (Firenze), 6: 43–68. Journal of the Entomological Society of South- Signoret, V. 1875. Essai sur les cochenilles ou ern Africa, 38: 125–130. gallinsectes (Homoptères – Coccides), 15e Ezzat, Y.M. & McConnell, H.S. 1956. A clas- partie. Annales de la Socété Entomologique de sification of the mealybug tribe Planococcini France (Serie 5), 5: 305–352. (Pseudococcidae: Homoptera). Bulletin of Tang, F.T. 1992. The Pseudococcidae of China. the Maryland Agriculture Experiment Station, Shanxi. 768 p. (In Chinese with English A-84: 1–108. summary). Ferris, G.F. 1950. Atlas of the Scale Insects of Tang, F.T. & Li, J. 1988. Observations on the North America. (Ser. 5. Vol. 5). The Pseudo- Coccoidea of Inner Mongolia in China. Inner coccidae (Part I). Palo Alto, California. 278 p. Mongolia University Press. 227 pp. Hambleton, E.J. 1935. Notes on Brazilian Tereznikova, E.M. 1975. Koktsidi. Chervetsi Pseudococcinae of economic importance plastinchasti, giganski ta boroshnisty [Cocci- with a description of four new species. Ar- nea. Ortheziidae, Margarodidae, Pseudococ- chivos do Instituto Biologico (São Paulo), 6: cidae]. Fauna Ukraini [Fauna of Ukraine], 105–120. 20(18). Kiev. 295 p. (In Ukrainian). Khadzhibeyli Z.K. 1983. Koktsidy subtropi- Ter-Grigorian, M.A. 1973. Chervetsi i shchitovki. cheskoi zony Gruzii [Coccids of the subtropi- Muchnistye chervetsi [Scale insects, mealy- cal zone of Georgia]. Tbilisi. 94 p. (In Rus- bugs (Coccoidea, Pseudococcidae)]. Fauna sian). Armyanskoy SSR [Fauna of Armenian SSR]. Kuwana, S.I. 1902. Coccidae (scale insects) of Erevan. 246 p. (In Russian). Japan. Proceedings of the California Academy Tranfaglia, A. & Tremblay, E. 1982. A morpho- of Sciences, 3: 43–98. logical comparison between Planococcus citri Marotta, S. 1990. Studio morfo-sistematico delle (Ris.), Planococcus ficus (Sign.) and their F1 specie italiane della famiglia Pseudococcidae hybrids (Homoptera: Coccoidea: Pseudococ- (Homoptera: Coccoidea). Ph.D. Thesis, Uni- cidae). Entomotaxonomia, 4: 1–5. versita degli studi di Napoli – Federico II, Umnov M.П. 1940. Karantinnye i drugie vidy Portici, Napoli. 104 p. koktsidy (Coccidae)Kryma [Quarantine and McKenzie, H.L. 1967. Mealybugs of California other pest species of scale insects (Coccidae) with taxonomy, biology, and control of North of the Crimea]. Simferopol. 64 p. American species (Homoptera: Coccoidea: Williams, D.J. 1985. Australian mealybugs. Lon- Pseudococcidae). Berkeley, California. 526 p. don. 431 p. Nasonov, N.V. 1908 (1909). On some new scale Williams, D.J. 2004. Mealybugs of Southern insects. Ezhegodnik Zoologicheskogo Muzeya Asia. Kuala Lumpur. 896 p. Imperatorskoi Akademii Nauk (St. Peters- Williams, D.J. & Moghaddam, M. 2000. Mealy- burg), 13: 471–499. (In Russian). bug species of the genus Planococcus Ferris Nedzelskii A. 1869. One of diseases of fruit plants in Iran (Homoptera: Coccoidea: Pseudococ- in the Crimea. Zemledelcheskaya gazeta (St. cidae) with a discussion of Planococcus vovae Petersburg), 19–23, 36–38. (In Russian). (Nasonov). Journal of Entomological Society Noyes, J.S. & Hayat, M. 1994. Oriental mealybug of Iran, 18(1/2): 32–43. parasitoids of the Anagyrini (Hymenoptera: Williams, D.J. & Watson, G.W. 1988. The Scale Encyrtidae). Wallingford, UK. viii+554 p. Insects of the Tropical South Pacific Region. Risso, A. 1813. Mémoire sur l’histoire naturelle Part 2. The Mealybugs (Pseudococcidae). des oranges, bigaradiers, limettiers, cédrat- London. 260 p. iers limoniers ou citroniers, cultivés dans le Received 12 May 2010 / Accepted 17 June 2010