Mealybugs of the Genus Rhizoecus Künckel D'herculais, 1878 (Homoptera: Pseudococcidae) of the Fauna of Russia and Adjacent Co

ZOOSYSTEMATICA ROSSICA, 18(2): 224–245 25 DECEMBER 2009

Mealybugs of the genus Rhizoecus Künckel d’Herculais, 1878 (Homoptera: Pseudococcidae) of the fauna of Russia and adjacent countries

E.M. DANZIG & I.A. GAVRILOV

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

A key and a review of 15 species inhabiting the territory of the former USSR are given. Rhizoecus microtubulatus Gavrilov & Danzig sp. nov. from Astrakhan is described. All discussed species are briefly morphologically described and illustrated. The lectotype is designated for Rh. vitis Borchsenius, 1949. Four new synonyms are established: Rhizoecus poltavae Laing, 1929 = Rh. desertus Ter-Grigorian, 1967 syn. nov. = Rh. pallidus Tereznikova, 1968 syn. nov.; Rh. tritici Borchsenius, 1949 = Rh. pratensis Borchsenius & Tereznikova, 1959 syn. nov.; Rh. albidus Goux, 1942 = Rh. gentianae Panis, 1968 syn. nov. Key words: scale insects, Homoptera, Pseudococcidae, Rhizoecus, new species

TAXONOMIC PART more) flagellate setae on dorsal side and usually with one apical setae on ventral side Genus Rhizoecus Künckel d’Herculais, 1878 of the same size as dorsal ones. Antennae with six or very rarely five broad, densely Rhizoecus Künckel d’Herculais, 1878: 163 settled segments. Legs short, with broad Rhizoecus: Hambleton, 1946: 50; 1976: 6; Borch- segments. Claw elongated. Coxae and tibiae senius, 1949: 174, Ferris, 1953: 426; McKen- without translucent pores. Ostioles poorly zie, 1967: 126; Danzig, 1980: 196; Ter-Grigo- visible, sometimes without trilocular pores rian, 1973: 89; Tereznikova, 1975: 246; Сox, 1987: 84; Tang, 1992: 53; Williams & Gran- and slender setae. Circulus usually present, ada de Willink, 1992: 494; Williams & Wat- more often conical and sclerotized, with cel- son, 1988: 213; Williams, 2004: 746; Kozár & lular pattern. Anal ring with 6 flagellate set- Benedicty, 2007: 126. ae and few distinctive large elongated pores. Ripersiella Tinsley, in Cockerell, 1899: 278 (type Multilocular pores usually present, located species Ripersia rumicis Maskell, 1892 by ori- on posterior abdominal sternites, rarely nu- ginal designation). merous on both body sides [Rh. poltavae á Ripersiella: Koz r & Benedicty, 2007: 380. Laing, 1929, Rh. brevipes (Goux, 1943)]. Pararhizoecus Goux, 1941: 197 (as subgenus; Trilocular pores numerous on both body type species Pararhizoecus petiti Goux, 1941 sides. Tubular ducts of different size and by monotypy). number. Peculiar bi- and tritubular ducts Type species: Rhizoecus falcifer Künckel are characteristic. Their length is usually d’Herculais, 1878 by monotypy. comparable to multilocular pores diameter. Diagnosis of the genus. Adult female. Sometimes tritubular pores can be signifi- Body elongate oval, often with almost par- cantly larger (Rh. vitis Borchsenius, 1949), allel margins, up to 2 mm long, covered and bitubular to the contrary smaller. Some with white powder wax; during the oviposi- species (for example, Rh. parvus Danzig, tion period females extract a loose wax sac. 1985) lack those ducts completely. Flagel- Anal lobes poorly developed, sometimes late and slender setae approximately equal with sclerotized plate, with two (sometimes on both body sides.

Mode of life. According to the mode of 1. Rhizoecus dianthi Green, 1926 life the majority of Rhizoecus species can (Fig. 1) be referred to soil insects; they live on thin roots of different plants, mostly perennial Rhizoecus dianthi Green, 1926: 175. Rhizoecus dianthi: Williams, 1962: 43; Cox, 1987: herbs, and only in rare cases inhabit over- 86; Kozár & Benedicty, 2007: 196; Danzig et head parts of host plants additionally. The al., 2008: 600. life cycle is typical for all mealybugs. Some Rhizoecus eluminatus McKenzie, 1960: 747 species are ovoviviparous laying eggs at the (USA: California). stage of germ band invagination or even Rhizoecus eluminatus: McKenzie, 1967: 388. with fully developed larvae inside (Tra- Rhizoecus pritchardi McKenzie, 1960: 749 (USA: peznikova & Gavrilov, 2008). At present, California). only five species are cytogenetically inves- Rhizoecus pritchardi: McKenzie, 1967: 400; Wil- liams & Nakahara, 1980: 336. tigated. Two species possess 2n=12, two 2n=10 and one 2n=8; all those five species Material. Five females from St. Petersburg, are characterized by a Lecanoid genetic several tens of series from Moscow: the Main Bo- system and bisexual reproduction (Nur et tanical Garden of the Russian Academy of Sci- ences, series of females from England (ZIN). al., 1987; Gavrilov, 2007; Gavrilov & Tra- Description. Adult female. Ostioles well- peznikova, unpubl.). developed. Circulus absent. Multilocular The genus is distributed all over the pores few, located only on three posterior world and comprises more than 200 spe- abdominal sternites. Tritubular ducts of cies. Due to the concealed mode of life and common size, few on dorsal body surface polyphagia some species have distributed and along margin of ventral body surface, together with the introduced host plants sometimes part of ducts accompanied by 1 far beyond their natural ranges. The fauna or 2 flagellate setae. Tubular ducts absent. of Russia and adjacent countries contains Systematical note. The species concerned 15 species of the genus, three of which are is probably identical to Rh. cyperalis (Ham- non-indigenous having been introduced belton, 1946) and Rh. nemoralis (Hambel- from North America and live only in green- ton, 1946) described from Central America, houses. In Western Europe there are some that was noted previously (Williams, 1962; more species of Rhizoecus. These are species Williams, Granada de Willink, 1992). from South France and Italy Rh. brevipes, Distribution. Reported only from green- Rh. brussieui (Goux, 1985), Rh. ovoides houses. Russia: St. Petersburg, Moscow. (Goux, 1943), Rh. periolanus (Goux, 1985), Europe, USA, Australia and New Zealand. Rh. petiti, Italian Rh. lelloi Mazzeo, 1995, Mode of life. Widely polyphagous on Rh. vidanoi Marotta & Tranfaglia, 1995 flowering plants. In St. Petersburg, it was and still not interpreted Rh. (?) targionii collected from Ophiopogon sp. (family Lili- (Cockerell, in Fernald, 1903), and also aceae). In Moscow, found on a range of plants two species described from Germany – Rh. in cactuses and succulents department. caesii Schmutterer, 1956 and Rh. franco- When the infestation is heavy the root sys- niae Schmutterer, 1956. tem and soil are entirely covered with white In greenhouses of Europe the follow- fluffy wax (Kozarzhevskaya, 1992). Serious ing species are also recorded according to pest of Saintpaulia spp. in California, cause the data of Jansen (2004): Rh. advenoides leaf shrinkage and plants death. Takagi & Kawai, 1971, Rh. aloes Williams & Pellizzari, 1997, Rh. americanus (Hamble- 2. Rhizoecus cacticans (Hambelton, 1946) ton, 1946), Rh. amorphophalli Betrem, 1940, (Fig. 2) Rh. elongatus Green, 1926, Rh. falcifer, Rh. hibisci Kawai & Takagi, 1971 and Rh. maas- Ripersiella cacticans Hambelton, 1946: 64 (Ec- bachi Jansen, 2003. uador).

Fig. 1. Rhizoecus dianthi, female, after Danzig et al. (2008), with specification.

Fig. 2. Rhizoecus cacticans, female, Moscow, greenhouse.

Ripersiella cacticans: Ferris, 1953: 432; Ham- Mode of life. Collected from Tanacetum belton, 1976: 17; Zahradnik, 1990: 41; Ko- vulgaris and Festuca sp. zarzhevskaya, 1992: 219; Kozár & Benedicty, 2007: 168. Rhizoecus inconspicuus Rhizoecus epiphylli Ferris, 1953: 442 (USA: Cali- 4. Danzig, 1971 fornia, Ontario). (Fig. 4) Rhizoecus epiphylli: McKenzie, 1960: 745 (= R. Rhizoecus inconspicuus Danzig, 1971: 372 (South cacticans). of Primorsk Terr.). Material. Large series from Moscow: the Rhizoecus inconspicuous: Danzig, 1980: 196. Main Botanical Garden of the Russian Academy Rhizoecus inconspicuus: Kozár & Benedicty, of Sciences, 14 specimens from St. Petersburg, 2007: 230. Minsk, Kiev, Donetsk and England (ZIN). Material. Holotype and paratypes (ZIN). Description. Adult female. Ostioles poor- Description. Adult female. Ostioles well- ly developed. Circulus conical, heavily scle- developed. Circulus conical. Anal lobes in rotized. Multilocular pores absent. Tritubu- holotype with a sclerotized plate with 3 lar ducts of common size, sparsed through- long and 1 or 2 shorter flagellate setae. Mul- out dorsal body surface, few along margin of tilocular pores few, located on three poste- ventral body surface. Tubular ducts narrow, rior abdominal segments. Tritubular ducts with parallel sides, few on both body sides. of common size, forming transverse rows on Distribution. Distributed throughout dorsal body surface, occur along the margin tropics and subtropics. In Russia and Eu- of ventral body surface. Tubular ducts short rope is common in greenhouses. and narrow, with parallel sides, numerous Mode of life. Polyphagous, live on differ- on both body sides. ent species of cactuses, harmful. Taxonomic Remarks. The close species Rh. advenoides Takagi & Kawai, 1971 also 3. Rhizoecus kazachstanus Matesova, 1980 connected with Artemisia spp. was de- (Fig. 3) scribed from Japan (Honshu). It differs in the absence anal lobes lacking a sclerotized Rhizoecus kazachstanus Matesova, 1980: 110 plate and minority of tubular ducts. Both (Kazakhstan). characters can result from interspecific or á Rhizoecus kazachstanus: Koz r & Benedicty, geographical variability, these species may 2007: 240. be identical. Material. Holotype and paratype (ZIN). Distribution. Russia: South of Primor- Description. Adult female. Ostioles poor- sky Terr. ly developed. Circulus conical. Anal lobes Mode of life. Females were collected in holotype with a sclerotized plate, in para- during oviposition on July 22 on Artemisia type and in material from Hungary the scle- sp. on weed-grown waste. rotized plate is not developed. Multilocular pores located on three posterior abdominal 5. Rhizoecus albidus (Goux, 1942) segments. Tritubular ducts of common size, (Fig. 5) numerous on both body sides. Tubular ducts dilated towards apex, located on both body Pararhizoecus albidus Goux, 1942: 40 (France). sides. Pararhizoecus albidus: Schmutterer, 1952: 394; Taxonomic Remarks. A figure based on Williams, 1962: 41; Tereznikova, 1975: 248; Danzig, 1980: 196; Kozár & Benedicty, the material from Hungary (Kozár & Bene- 2007: 140. dicty, 2007) differs from the type material Rhizoecus uniporus Borchsenius, Tereznikova, in a larger number of tubular ducts. 1959: 323 (Ukraine). Distribution. Western and northern Ka- Rhizoecus uniporus: Tereznikova, 1975: 248 (= R. zakhstan. Hungary. albidus); Ter-Grigorian, 1973: 92.

Fig. 3. Rhizoecus kazachstanus, female, holotype.

Fig. 4. Rhizoecus inconspicuus, female, after Danzig (1971), with additions.

Fig. 5. Rhizoecus albidus, female, after Danzig (1980), with specification.

Rhizoecus gentianae Panis, 1968 (France, Italy), lar pores, located on both body sides, not syn. nov. numerous. Tubular ducts wide, located on Rhizoecus gentianae: Kozár & Benedicty, 2007. both body sides. Material. Paratype and 19 females from Rus- Distribution. Known only from the type sia (Irkutsk Prov., South of Primorsk Terr.), series. Ukraine, Armenia, Switzerland, England and the Mode of life. Collected from roots of Iran (ZIN). vine. Description. Adult female. Ostioles well- developed. Circulus conical. Multilocular 7. Rhizoecus mexicanus (Hambelton, 1946) pores absent. Tritubular ducts of common (Fig. 7) size, forming row along dorsal body surface margin and middle line, single pores occur Ripersiella mexicana Hambelton, 1946: 67 (Mex- in middle part of body; on ventral body sur- ico, imported from California). face arranged only along body margin. Tu- Ripersiella mexicana: Hambelton, 1976: 37; Kozár & Benedicty, 2007: 500. bular ducts short and narrow, with parallel Rhizoecus mexicanus: Hambleton, 1976: 37; sides, few. Williams & Granada de Willink, 1992: 549; According to the description and figure Gavrilov, 2004: 115. of Panis (1968) and also comments of Kozár Material. One paratype and 4 females from & Benedicty (2007), Rhizoecus gentianae Russia and the Netherlands (ZIN). differs from Rh. albidus in longer labium Description. Adult female. Ostioles poor- only. This character varies individually and ly developed. Circulus slightly convex. Mul- can not be used for the species separation. tilocular pores located on two posterior ab- Distribution. Russia: Irkutsk Prov. and dominal sternites and penultimate abdomi- South of Primorsky Terr.; Ukraine, Arme- nal tergite. Bitubular ducts close in size to nia. Europe: widely, Iran. multilocular pores, occur throughout dorsal Mode of life. Live mainly on gramineous body surface and along margin of ventral plants; in the Far East females were collect- body surface, single ducts located in middle ed in the end of August. part of posterior abdominal and thoracical sternites. Tubular ducts of middle size, with 6. Rhizoecus vitis Borchsenius, 1949 parallel sides, sclerotized apex, numerous on (Fig. 6) both body sides. Distribution. Greenhouses of Russia Rhizoecus vitis Borchsenius, 1949: 175 (Ukraine, (St. Petersburg) and the Netherlands, Crimea). Rhizoecus vitis: Tereznikova, 1975: 246; Kozár & Mexico, USA. Benedicty, 2007: 252. Mode of life. Live on cactuses; in St. Material. Lectotype (designated here). ZIN, Petersburg and the Netherlands collected slide № 7–40; female, Ukraine, Crimea, Balak- from Opuntia spp. lava Reg., 15 July 1939, coll. Sheffer. Paralecto- types. Six females, same data as lectotype, and 6 8. Rhizoecus poltavae Laing, 1929 females from different regions of Crimea. (Fig. 8) Description. Adult female. Differs from the other species in 5-segmented antennae. Rhizoecus poltavae Laing, 1929: 469. Ostioles well-developed. Circuli 3 in num- Rhizoecus poltavae: Borchsenius, 1949: 177; Ham- ber, large, flat. Anal lobes with large, convex belton, 1946: 50; Tereznikova, 1975: 251. Ripersiella poltavae: Kozár & Benedicty, 2007: sclerotized plate with 11–15 long flagellate 524. setae. Multilocular pores numerous on two Rhizoecus desertus Ter-Grigorian, 1967: 93 (Ar- posterior abdominal sternites. Tritubular menia), syn. n. ducts larger than in other species of Russian Rhizoecus pallidus Tereznikova, 1968: 377 (Ukra- fauna, significantly larger than multilocu- ine), syn. n.

Fig. 6. Rhizoecus vitis, female, lectotype.

Fig. 7. Rhizoecus mexicanus, female, after Gavrilov (2004).

Fig. 8. Rhizoecus poltavae, female, lectotype.

Lectotype of Rh. poltavae (designated cies Goux points to the absence of bitubular here). ZIN, female, Ukraine, “Rhizoecus pol- ducts. When studying the paratype of this tavae Laing, Poltava, Ogloblin”. Slide with- species we also failed to reveal it. Tubular out number, specimen in black circle. ducts in Rh. brevipes and Rh. poltavae also Paralectotypes. ZIN; 4 more females to- differ in it’s structure. gether with lectotype, 6 females on two oth- In the description and figure made by er slides with the same label and 4 females Tereznikova (1975) any bitubular ducts from the same locality but with the date are missing. We examined the material on 28.04.1925. In the species description an er- which the description and the figure were ror was made in the type locality indication based. “Crimea: Poltava”, in London (the Natural Distribution. Russia: Krasnodar Terr. History Museum) four slides with the label (Anapa Reg.); Ukraine, Armenia, western Rh. poltavae Laing are deposited, on two Kazakhstan, Kirghizia. Italy. of them also “Crimea: Poltava” is given, on Mode of life. Host plants for the type se- the others “Odessa, Ukraina, Poltava”. On ries of Rh. poltavae are unknown; Rh. deser- every slide the date 28.04.1925 and the col- tus was described from Veronica armena, Rh. lector A.N. Kiritshenko are indicated. It is pallidus – from a gramineous plant; in Ka- evident that it is the same material that was zakhstan it was collected from Spiraea sp. collected by Ogloblin in Poltava and sent to London by Kiritshenko. For that reason we 9. Rhizoecus tritici (Borchsenius, 1949) consider the English material as belonging (Fig. 9) to the type series and, thus, paralectotypes. Other material examined. Holotype of Rhizoecus tritici Borchsenius, 1949: 177 (Russia: Rh. desertus and paratype of Rh. pallidus, Orenburg Reg.). Rhizoecus tritici: Tereznikova, 1975: 251. large series of females and nymphs from Ripersiella? tritici: Kozár & Benedicty, 2007: 564. western Kazakhstan, one female from Kir- Rhizoecus pratensis Borchsenius & Tereznikova, ghizia (Kungey-Alatau). 1959: 322 (Ukraine: Transcarpathia), syn. n. Description. Adult female. Ostioles not Rhizoecus pratensis: Tereznikova, 1975: 248 (as developed. Circulus slightly convex. Mul- Ph. halophilus). tilocular pores numerous on both body sides. Ripersiella pratensis: Kozár & Benedicty, 2007: 526. Bitubular ducts smaller than multilocular Material. Holotypes (monotypes) of Rh. pores, located on dorsal body surface and pratensis and Rh. tritici and a female series from along margin of ventral body surface, single Kazakhstan. ducts located in middle part of posterior Description. Adult female. Ostioles well- abdominal and thoracic sternites. Tubular developed. Circulus not slightly convex. ducts wide, with a small collar at the base, Multilocular pores arranged on three pos- occur on both body sides, few. terior abdominal sternites. Bitubular ducts Taxonomic Remarks. The synonymy is smaller than multilocular pores, few on both established while comparing of the types of body sides. Tubular ducts wide, with paral- Rh. poltavae, Rh. desertus and Rh. pallidus. lel sides, sclerotized at base, numerous on Rh. lelloi Mazzeo, 1995, that was described both body sides. from Italy and noted for Hungary, and also Systematical notes. Synonymy was estab- Rh. caesii Schmutterer, 1956 from Germa- lished when comparing of the holotypes of ny, are probably identical to Rh. poltavae. Rh. tritici and Rh. pratensis. In the primary Unfortunately, we had no opportunity to description and in the book by Tereznikova become acquainted with the types of those (1975) bitubular ducts in Rh. pratensis are species. To this species group with numer- missing. Rhizoecus petiti is close to Rh. pra- ous multilocular pores Rh. brevipes Goux, tensis, differs in character of tubular and tri- 1943 also belongs. In description of this spe- tubular ducts.

Fig. 9. Rhizoecus tritici, female, holotype.

Distribution. Russia: Orenburg Reg.; body surface. Tubular ducts small, with par- western Kazakhstan, Ukraine. Italy. allel sides, numerous on both body sides. Mode of life. Collected from Festuca sul- Distribution. Russia: Karelia. Europe: cata and Koeleria gracilis. widely. Mode of life. Polyphagous species. In 10. Rhizoecus ornatus Borchsenius, 1949 Karelia collected in littoral on Triticum (Fig. 10) vulgaris, Leumus arenarius, Elytrigia, Plan- tago maritima, Trifolium vulgaris. Rhizoecus ornatus Borchsenius, 1949: 176 (Ar- menia) 12. Rhizoecus microtubulatus Gavrilov & Rhizoecus ornatus: Ter-Grigorian, 1973: 90; Danzig, sp. nov. Kozár & Benedicty, 2007: 288. Rhizoecus ornatoides Tang, 1992: 62 (a substitute (Fig. 12) name erroneously introduced for Rhizoecus Holotype. ZIN, K No 407; female, Russia, ornatus Borchsenius, 1949, a senior hom- Astrakhan, semi-desert near Tinaki Lake, on thin onym of Rh. ornatus Hambelton, 1976; the roots of Artemisia vulgaris, 13 May 2004, coll. latter species is now in Benedictycoccina á I.A. Gavrilov. Koz r & Foldi, 2004). Paratype. ZIN; female, same data as holotype. Material. The species known from the holo- Description. Adult female. Ostioles poor- type (ZIN). ly developed. Circulus conical. Multilocular Description. Adult female. Ostioles well- pores located on three posterior abdominal developed. Circulus conical. Anal lobes with sternites. Bitubular ducts very small, com- large, flat, sclerotized plate with 3 long and parable to trilocular pores in size. Basing on 1 short flagellate setae. Multilocular pores the available material, it is hard to judge the located on two or three posterior abdomi- location of bitubular and tritubular ducts. nal sternites. Bitubular ducts (large trilocu- They seem to form transverse rows on ven- lar in description by Borchsenius and Ter- tral body surface. Tubular ducts wide, dilat- Grigorian, trilocular in Kozár & Benedicty), ing to strongly sclerotized apex. equal to multilocular pores in size, forming Comparative Remarks. The new species transverse rows on dorsal body surface, sin- differs from the other Palaearctic species gle on ventral body surface. Tubular ducts of the genus Rhizoecus in structure of the small, with parallel sides and sclerotized bitubular ducts of a very small size (see apex, present on both body sides. Fig. 12); their size is similar to the size of Distribution. Armenia. trilocular pores. Mode of life. Collected under a stone. Etymology. Species name is derived from the main diagnostic character of the new 11. Rhizoecus halophilus (Hardy, 1868) species, small size of the bitubular ducts. (Fig. 11) Mode of life. Collected in the beginning of oviposition period. Coccus halophilus Hardy, 1868: 136 (Scotland). Rhizoecus halophilus: Williams, 1962: 47. 13. Rhizoecus parvus Danzig, 1985 Ripersiella halophila: Kozár & Benedicty, 2007: 458. (Fig. 13) Material. 14 females from Karelia, 2 from Italy, 1 from Ireland, female series collected by I.A. Gavr- Rhizoecus parvus Danzig, 1985: 121 (Russia: ilov in Bulgaria (Rila Mountains) (ZIN). Northern Caucasus). á Description. Adult female. Ostioles well- Rhizoecus parvus: Koz r & Benedicty, 2007: 511. developed. Circulus conical. Multilocular Material. Holotype and paratypes (ZIN). pores located on three posterior abdominal Description. Adult female. Ostioles poor- sternites. Bitubular ducts equal to multiloc- ly developed. Circulus flat. Multilocular ular pores in size, few, located only on dorsal pores located on V–VII abdominal terg-

Fig. 10. Rhizoecus ornatus, female, holotype.

Fig. 11. Rhizoecus halophilus, female, Russia: Karelia.

Fig. 12. Rhizoecus microtubulatus, sp. nov., female.

Fig. 13. Rhizoecus parvus, female, holotype.

© 2009 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 18(2): 224–245 E.M. DANZIG & I.A. GAVRILOV. MEALYBUGS OF THE GENUS RHIZOECUS OF RUSSIA 243 ites and IV–VIII abdominal sternites. Bi- 23(16). Bitubular ducts small, in size comparable and tritubular ducts not found, irregular with trilocular pores (Fig. 12). Tubular ducts strongly sclerotized pores present. Tubular broad, dilating towards heavily sclerotized ducts dilating to strongly sclerotized apex, apex ...... Rh. microtubulatus sp. nov. numerous on both body sides. 24 (7). Tritubular and bitubular ducts absent, sclerotized irregular pores with obscure Distribution. Known from the type local- structure present. Tubular ducts short, with ity only. collar at the base ...... Rh. parvus Mode of life. Collected from Sempervi- vum caucasicus and Artemisia vulgaris. ACKNOWLEDGEMENTS

KEY TO SPECIES The authors are grateful to Dr. Danièle Mat- ile-Ferrero (Museum National d’Histoire Na- 1(6). Occur in greenhouses only. 2(5). Tritubular ducts present, bitubular ducts turelle, Paris) for the opportunity to work with absent. the types of species, described by L. Goux, and to 3(4). Multilocular pores present. Circulus and Jon Martin (British Museum of Natural History, tubular ducts absent ...... Rh. dianthi London) for clarifying the label data of the type 4(3). Multilocular pores absent. Circulus and tu- material of Rh. poltavae. The work was supported bular ducts present ...... Rh. cacticans by Russian Foundation for Basic Research (proj- 5(2). Tritubular ducts absent, bitubular ducts ects Nos 08-04-00787 and 09-04-91229-Ста) present ...... Rh. mexicanus and by a Special grant from the Government of 6(1). Occur in open ground conditions. St. Petersburg (for IG). 7(24). Tritubular or bitubular ducts present, sclerotized irregular pores absent. Tubular REFERENCES ducts of different types. 8(15). Tritubular ducts present. Ben-Dov, Y. 1994. A systematic catalogue of the 9(14). Single circulus present. Antennae 6-seg- mealybugs of the World (Insecta: Homoptera: mented. Coccoidea: Pseudococcoidae and Putoidae). 10(13). Multilocular pores present. With data on geographical distribution, host 11(12). Tubular ducts broad, dilating towards apex. Anal plate not always visible ...... plants, biology and economic importance. In- ...... Rh. kazachstanus tercept Limited, Andover. 686 p. 12(11). Tubular ducts narrow, with parallel sides Borchsenius, N.S. 1949. Coccoidea. The Fam- ...... Rh. inconspicuus ily Pseudococcidae. Fauna SSSR, Nasekomye 13(10). Multilocular pores absent . . .Rh. albidus khobotnye (The Fauna of the USSR, Ho- 14(9). 3 circuli present. Antennae 5-segmented . moptera–Heteroptera), 7. Moscow. 383 p...... Rh. vitis (In Russian). 15(8). Bitubular ducts present, sometimes few. Borchsenius, N.S. & Tereznikova, E.M. 1959. 16(23). Bitubular ducts of common size; their Two new species of mealy bugs of the ge- length equal or close to diameter of multiloc- nus Heliococcus Šulc (Insecta, Homoptera, ular pores. Tubular ducts narrow or broad, Coccoidea). Zoologicheskiy Zhurnal, 38(3): with parallel sides. 307–321. 17(18). Multilocular pores numerous on both Cockerell, T.D.A. 1899. Tables for the determi- body sides ...... Rh. poltavae nation of the genera of Coccidae. Canadian 18(17). Multilocular pores located only on last Entomologist, 31: 273–279. abdominal segments. Cox, J.M. 1987. Pseudococcidae (Insecta: He- 19(20). Anal plate well-developed, sclerotized . . mi ptera). Fauna of New Zealand, 11. Wel- ...... Rh. ornatus lington. 229 p. 20(19). Anal plate not developed. Danzig, E.M. 1980. Coccoids of the Far East 21(22). Tubular ducts broad, with collar at the USSR (Homoptera, Coccinea) with phylo- base. Circulus slightly convex . . . . .Rh. tritici genetic analysis of scale insects fauna of the 22(21). Tubular ducts narrow, without collar at world. Opredeliteli po faune SSSR (Keys to the base. Circulus strongly convex ...... the Fauna of the USSR), 24. Nauka, Lenin- ...... Rh. halophilus grad. 368 p. (In Russian).

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