ZOOSYSTEMATICA ROSSICA, 24(2): 236-259 25 DECEMBER 2015 System of generic groups in mealybugs (Homoptera: Coccinea: Pseudococcidae) Система групп родов мучнистых червецов (Homoptera: Coccinea: Pseudococcidae) I.A. GAVRILOV-ZIMIN И.А. ГАВРИЛОВ-ЗИМИН Zoological Institute, Russian Academy of Sciences, 1 Universitetskaya Emb., St Petersburg 199034, Russia. E-mail: [email protected] The paper provides a brief conspectus of the system of morphological generic groups, elaborated ear- lier by the author basing on the total taxonomic revision of Palaearctic mealybugs. Here the system is complemented by the analysis of all 249 genera of the world fauna. Borders of two generic groups are reconsidered and two else groups (with mainly Oriental and Australasian genera) are included in the system. Main taxonomic characters of generic rank are discussed and illustrated. Статья представляет собой краткий конспект системы морфологических групп родов мучнистых червецов, предложенной автором ранее на основе тотальной ревизии палеарктической фауны. В настоящей работе эта система дополнена обзором всех 249 родов мировой фауны псевдококцид. Объем двух родовых групп пересмотрен; еще две группы, включающие преимущественно рода ориентальной и австралазийской фаун добавлены в систему. Рассмотрены и проиллюстрированы основные таксономические признаки родового уровня. Key words: scale insects, mealybugs, systematics, morphology, ovoviviparity Ключевые слова: кокциды, мучнистые червецы, систематика, морфология, яйцеживо- рождение INTRODUCTION parative analysis of the basic morphologi- cal characters of mealybugs, discussed in The Pseudococcidae or mealybugs are detail all main modern contradictions in one of two largest scale insect families and the higher classification of the family and comprises more than 2000 species in world introduced a system of 19 morphological fauna. Unfortunately, general classifica- generic groups, most of which seem to tion and phylogeny reconstruction for the be monophyletic (holophyletic or para- family have been poorly elaborated till now phyletic) and will be probably considered in view of the numerous problems with in future as taxa of the formal nomen- the correct, unequivocal evolutional inter- clatural rank (for example, tribes). Some pretation of the mealybug morphological other morphological groups are perhaps (and some other) characters as plesiomor- polyphyletic and combine genera which phic, apomorphic, convergent or parallel demonstrate convergent similarity only; (homoplasy). In our recent total review of however, the decision of this problem is Palaearctic mealybugs (Danzig & Gavrilov- impossible basing on the present basis of Zimin, 2014; 2015) we have provided com- our knowledge. © 2015 Zoological institute, Russian Academy of Sciences I.A. GAVRILOV-ZIMIN. SYSTEM OF GENERIC GROUPS IN MEALYBUGS 237 Here I am providing a synopsis of the DISCUSSION elaborated system, a discussion of some ad- The main taxonomic characters used ditions to the system, basing on analysis of in the analysis of the genera (Fig. 1) non-Palaearctic genera and a conspectus of all mealybug genera of the world fauna [authorship of the generic names see below in with preliminary grouping of them accord- the system of generic groups] ing to the system. Two generic groups used Wax glands (Table). The structure and in the revision of Palaearctic fauna (Danzig distribution of wax glands are the main di- & Gavrilov-Zimin, 2014, 2015), g/g Trabu- agnostic characters in the systematics of tina and g/g Planococcus, are included here mealybugs and all scale insects. Cuticular correspondingly to g/g Nipaecoccus and sclerotized parts of these glands are clearly g/g Trionymus (see below). On the other visible because they absorb stain during hand, two other groups, which combine preparation of scale insects and can be sub- mainly Oriental and Australasian genera, divided into 3 main groups: discoidal glands, g/g Pedronia and g/g Allomyrmococcus are cylindrical glands and glands connected added to the system. with conical setae (“spines” in non-English The analysis and discussion below are literature). {In the English coccidological based on the study of the type species of tradition the term “wax glands” is usually 249 genera, most of which are represented not used, probably because the soft parts of (as type and non-type specimens) in the the glands are lost during preparation and collections of Zoological Institute, Rus- only “ducts” and “pores” are visible on spec- sian Academy of Sciences (St Petersburg) imens mounted in Canada-balsam. In Rus- and Muséum National d’Histoire Naturelle sian and other old European coccidological (Paris). Some material was also loaned by schools the term “glands” is widely used, be- me from the collection of the British Mu- cause there are no special terms to indicate seum of Natural History (London) and the soft parts of the wax glands.} Plant Protection Institute of Hungarian Discoidal glands produce powdery wax Academy of Sciences (Budapest). In addi- and can be subdivided in the following sub- tion to the study of the materal I used for types. the analysis the morphological descriptions 1) Simple discoidal pores; have a single and illustrations published in the compre- opening (loculus). Such pores are often hensive faunistic reviews by following au- scattered in more or less quantity on both thors for the main non-Palaearctic zoogeo- body surfaces of mealybugs and/or present graphical regions of the world: for Nearctic in groups with other wax glands or associat- (Ferris, 1950, 1953; McKenzie, 1967), for ed with different types of setae. Monotypic Neotropical (Williams & Granara de Wil- genus Hopefoldia is characterized by very link, 1992), for Oriental (Williams, 2004) peculiar compact groups of discoidal pores, and for Australasian (Ferris, 1948; Beards- so-called “poraria” located on sclerotized ley, 1964, 1966, 1971; Williams, 1985; Cox, plates on both body sides. 1987; Williams & Watson, 1988). The Afro- 2) Trilocular (3-locular) pores have 3 tropical region has not been covered by the excretory openings and approximately tri- general taxonomic revision till now, but the angular in form. These pores are one of the following basic publications help to unders- synapomorphic characters of mealybugs and tand the main morphological trends in the the monotypic family Phenacoleachiidae evolution of the mealybugs of the region: and can be found in majority of mealybugs Mamet, 1940, 1951, 1954, 1962, 1967; De on all body surface. Only some mealybug Lotto, 1957, 1958, 1964, 1967, 1969, 1977; genera, such as from g/g Heterococcus, g/g Millar, 2002). Boreococcus, g/g Neotrionymus, g/g Metad- © 2015 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 24(2): 236–259 238 I.A. GAVRILOV-ZIMIN. SYSTEM OF GENERIC GROUPS IN MEALYBUGS Quinquelocular pore Eye Claw digitules Anterior ostiolus Claw with denticle Anterior spiracle Trilocular pore Simple pore Conical seta Circuli Flagellate setae Simple tubular duct Pores Spinulae Anus Multilocular pore Cerarius C Vulva 18 Setae of anal apparatus Fig. 1. Main taxonomic characters of mealybug females (Fonscolombia tomlinii (Newstead, 1892), Russia: Voronezh Prov.). © 2015 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 24(2): 236–259 I.A. GAVRILOV-ZIMIN. SYSTEM OF GENERIC GROUPS IN MEALYBUGS 239 enopus, and also some genera from other Cylindrical glands (Table 1) produce groups demonstrate reduction in the num- wax filaments. There is a great diversity of ber of trilocular pores or their secondary these glands, probably, exceeding the diver- total absence. Sometimes trilocular pores sity of such glands in other scale insects all are scattered on body very irregularly; thus, together. they can be absent or very few in medial 1) Simple tubular ducts. They have one zone of the thoracic and anterior abdominal simple excretory duct. The opening of this sternites (as, for example, in some species of duct usually lies on the surface of the cu- the species-rich genus Phenacoccus) or form ticle or more rarely it is slightly raised or dense marginal or medial bands (as in Trim- slightly invaginated from the surrounded erococcus and Kiritshenkella, respectively). cuticle area. Most genera of subfamily Phe- The size of the trilocular pores is usually nacoccinae have only this type of cylindri- equal throughout the body. However, in the cal glands. The deviation of this type is so- species of g/g Puto, dorsal trilocular pores called “bottle-shaped” ducts in the species (and especially cerarian ones) are larger of g/g Coccidohystrix: distal (distant from than ventral pores. Trilocular pores of pe- the excretory opening) part of such duct is culiar structure are known in some species enlarged in the form of a bottle. and genera, for example, in Cucullococcus 2) Collar tubular ducts. In these glands arrabidensis (Neves, 1954) (pores without the duct is surrounded by collars of differ- fringe) and in the species of Iberococcus ent sizes and forms. The greatest diversity (pores are strongly convex and located in a of collar ducts takes place in the subfam- deep cup). ily Pseudococcinae, whereas among more 3) Quinquelocular (5-locular) pores primitive Phenacoccinae these ducts are have one central loculus and 5 peripheral known in several genera only, for example loculi; sometimes central loculus absent. in Heliococcus, Seyneria, Ehrhornia. There These pores are present mainly in the species are at least the following main variants of of the subfamily Phenacoccinae (in Pseu- collar ducts. dococcinae they are known mainly in g/g a) Fungiform ducts. The collar of these Metadenopus) and usually scattered on the ducts is very wide and similar to a cap of ventral surface of body, especially around fungus. Most of species with fungiform mouthparts and near spiracles. In some cas- ducts are members of g/g Pseudococcus es (for example, in g/g Heterococcus and g/g and only some species with such ducts are Metadenopus) 5-locular pores are numerous known in other genera, for example, in the on both body sides and replace partly or to- genera Mirococcopsis and Humococcus. tally reduced trilocular pores. b) Oral collar ducts, with narrow collar 4) Multilocular pores usually have one which surrounds the excretory opening and central loculus and 8–10 peripheral loculi part or all tube of the duct. Such ducts are and form transverse rows on abdominal most common and known in most genera of sternites. More rarely multilocular pores the subfamily Pseudococcinae. The length present on other surface of body and/or of the collar can be rather different, from show a peculiar distribution, structure and very small to longer than the length of the size. For example, in Mirococcus sphaeroides duct itself. Danzig, 1975 some of multilocular pores c) Middle collar ducts, with collar lo- have several central loculi; species of the cated in the middle part of the duct (for genera Pelionella and Rosebankia show 2 example, in Volvicoccus volvifer Goux, different types of multilocular pores in the 1945, Metadenopus caudatus (Borchsenius, same female. In g/g Neotrionymus, multiloc- 1958), etc.). ular pores replace partly or totally reduced d) Double collar ducts. Proximal part of trilocular pores. such collar is always more sclerotized than © 2015 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 24(2): 236–259 240 I.A. GAVRILOV-ZIMIN. SYSTEM OF GENERIC GROUPS IN MEALYBUGS Table 1. Wax glands in Pseudococcidae. Discoidal wax gland s Simple discoidal pore s Nudicauda nigra Trilocular pore s Cucullococcus Iberococcus Quinquelocular por es Multilocular pore s Pelionella Cylindrical wax glands Simple tubular ducts Coccidohystrix Fungiform ducts Oral collar ducts Middle collar ducts Collar tubular ducts Double collar ducts Crateriform duct s Heliococcus Mitral duct s Ehrhornia, Discococcus Bitubular /tritubular ducts (in g/g Rhizoecus) Peculiar types Nudicauda Ferrisia © 2015 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 24(2): 236–259 I.A. GAVRILOV-ZIMIN. SYSTEM OF GENERIC GROUPS IN MEALYBUGS 241 the distal part (for example, in some Triony- along body margin, cerarii, which is one of mus, Paraputo, etc.). unique apomorphic characters of Pseudo- f) Crateriform ducts. The collar of such coccidae. Structure and number of cerarii ducts bears one or several conical setae. are important taxonomic characters which These are known in the genus Heliococcus are used in the classification and identifica- only. tion at the species level and at higher taxa g) Mitral ducts; resemble a headwear level. The number of cerarii usually ranges (“mitre”) of orthodox religious superiors. from 1 to 18 pairs (first pair on head and These ducts are known, for example, in the the last pair on last abdominal tergite) and genera Ehrhornia, Discococcus, Paradisco- their exact number is usually stable inside coccus. of the species. More rarely there are small 3) Bitubular and tritubular ducts are individual variations of this number. An structures, consisted of 2 or 3 crossed ducts, extraordinary variability of the number is with or without general collar; these are currently known in some species only. For present in many (but not in all!) species of example, Dysmicoccus angustifrons (Hall, g/g Rhizoecus. 1926) has from 2 to 10 pairs of cerarii, Atro- 4) Peculiar types of tubular ducts are coccus alhagii (Hall, 1926) – from 2 to 15 known in different genera. For example, in pairs. Evolutional and individual reduction the species of the genus Nudicauda, and in in the number of cerarii always occurs in the the monotypic genus Malekoccus, the duct following succession: thoracic and anterior opening is encircled by a flat sclerotized abdominal cerarii become absent initially; ring; in the species of pantropical genus Fer- followed by the cerarii on head, and at the risia, the opening is encircled by a sclero- end the anal pair (С ) is reduced. 18 tized zone of the cuticle with several associ- In some mealybugs the normal cerarii ated flagellate setae. are totally absent or in the place of the last In addition to these rather large ducts cerarian pair there are one or several flag- which are well visible under usual trans- ellate setae, accompanied by group of tri- lucent microscope, many mealybugs have locular pores. We consider such structures numerous poorly visible minute microtubu- as pseudocerarii. Numerous pseudocerarii, lar ducts, which can be located on different distributed along all body margin are a very parts of the body and sometimes on the hind rare character which is known in monotypic coxae. Fine structure and function of these Madagasia and Telocorys and in Dysmicoc- microducts are unknown till now. cus trispinosus (Hall, 1923), which is prob- Conical setae have different forms and ably must be transferred to Telocorys. sizes (Fig. 2) and produce consistent wax. In g/g Puto and g/g Paraputo, the mar- Especially large and numerous conical setae ginal cerarii are more numerous, up to 26 are known in the species of g/g Puto, g/g pairs. Moreover, additional (non-marginal) Coccidohystrix and g/g Pedronia. In con- cerarii can present in the in medial zone of trast, in some desert mealybugs, conical se- dorsum. In the monotypic genera Trimero- tae are totally lost and the body is covered coccus and Eastia and in many species of g/g by flagellate and/or hair-like setae only. Paraputo the marginal cerarii are fused in a In most of mealybugs the conical setae solid marginal band. have pointed or slightly roundish apex and In some species of Coccidohystrix, normal only in the genera Rastrococcus and Stemma- cerarii are replaced by groups of large coni- tomerinx the conical setae are sharply trun- cal setae on cuticular tubercles, without tri- cated (see Fig. 2). The conical setae with locular or 5-locular pores – mamelons. associated discoidal glands (usually with Flagellate and hair-like setae of differ- trilocular pores, more rarely with 5-locu- ent size are more or less numerous on ven- lar pores) form multiple binate structures tral body side of all species. Dorsal flagellate © 2015 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 24(2): 236–259 242 I.A. GAVRILOV-ZIMIN. SYSTEM OF GENERIC GROUPS IN MEALYBUGS and/or hair-like setae are present mainly Fig. 3. The compound structure of the anal in the species of subfam. Pseudococcinae. apparatus is connected with the mode of Amongst Phenacoccinae these setae (usual- mealybug nourishment. Constant sucking of ly together with conical setae) are present, sweet sap from the plant floem leads to prob- for example, in Palaearctic genera Fonscol- lems with eliminating the surplus of water ombia and Perystrix. and sugar (honeydew) from the organism. Ostioles represented by slit-like open- The honeydew would easily cover the anal ings, located symmetrically as one or 2 pairs segment and occlude the anus if lacking spe- on dorsal (rarely on margin of ventral) sur- cial morphological structures that envelop face of body. These structures are one of the honeydew drop with wax in order to the unique synapomorphic characters of discard it from the body. Secondary simpli- mealybugs (including “Putoidae” of some fication of the anal apparatus can be con- authors) and the monotypic family Phen- nected with the feeding of sap directly from acoleachiidae. The position of the ostioles the plant cells or by obligate symbiosis with is constant; anterior pair always located on ants which feed on the excreted honeydew. anterior part of cephalothorax and the pos- Legs usually small in comparison with terior pair – on VI or VII abdominal tergites body; in the species of g/g Antonina legs or between them. Margins (lips) of ostioles significantly reduced or totally absent. usually slightly raised above surrounded Tarsus 1-segmented, with 2 thin setae at cuticle and bearing the same cuticular the apex, so-called “tarsal digitules”. Claw structures, i.e., different setae and pores. In with or without a denticle; this is impor- the Oriental mealybugs of g/g Allomyrmo- tant taxonomic character that is tradition- coccus, the ostioles are highly convex and ally used for the separation of two subfami- strongly sclerotized. In very rare cases, the lies of Pseudococcidae. Claw digitules are ostioles are secondary absent though they usually slightly longer than claw and have are known in related species and genera or a clavate apex; very rarely they are short even in the larvae of the same species. and/or pointed. In the species of g/g Allo- Anal apparatus of a compound struc- myrmococcus claw digitules are widely ex- ture: flat sclerotized ring around anus, one panded. All legs more or less equal in size, inner row of round or oval pores, one outer but in Mongolian endemic, Mirococcus fos- row of spinulae and 6 setae, which are usu- sor Danzig, 1983, anterior legs of burrowing ally slightly longer than the diameter of anal type: femur, tibia and tarsus strongly en- ring. Such most common type of anal appa- larged and claw elongated, with 2 denticles; ratus we name as complete apparatus (Fig. claws of other legs of this species are also 3). In some species the number of spinulae elongated, but without denticles. Claws is more numerous – 2–4 rows; such type of with 2 denticles are also known in some anal apparatus we consider as complicated. other mealybugs, for example, in Perystrix On the contrary, in some species, especially ulmaria Gavrilov, 2004; the second denticle in deserts, anal apparatus is strongly sim- is always smaller than the first one. Unusu- plified, with reduced number of pores and ally enlarged legs are known in the species spinulae or even with their total absence. of Australian Pseudoripesia and South Af- Additionally, many species have more or less rican Diversicrus. Hind coxae (more rarely developed anal tube, invaginated inside of hind femurs and tibiae) of many species, anal segment of body. In most species of g/g mainly from the subfamily Pseudococcinae, Rhizoecus, pores and spinulae of anal appa- bearing so-called “translucent pores” or, ratus are enlarged and irregular-shaped. In more rarely, very small microtubular ducts. the genera Rhodania and Pararhodania, anal In the species of g/g Antonina numerous ring strongly enlarged and sclerotized. Dif- and closely spaced translucent pores or mi- ferent types of anal apparatus are shown on crotubular ducts form two compact groups © 2015 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 24(2): 236–259 I.A. GAVRILOV-ZIMIN. SYSTEM OF GENERIC GROUPS IN MEALYBUGS 243 Conical setae Flagellate setae Hair-like setae Cerarius with multiple Usual cerarius Pseudocerarius Mamelon setae on sclerotized plate Fig. 2. Setae, cerarii and cerarius-like structures. on metathorax and anterior abdominal ster- cies in nature is higher. Some species-rich, nites instead of lacked hind legs. but morphologically heterogenous genera Ovoviviparity. We (Gavrilov-Zimin & such as Phenacoccus, Pseudococcus, Triony- Danzig, 2012; Danzig & Gavrilov-Zimin, mus include oviparous and obligate ovovi- 2014) consider ovoviviparity as a plesio- viparous species and also species with facul- morphic character of all Pseudococcidae. tative ovoviviparity. This character was probably inherited by mealybugs from the obligate ovoviviparous Family Pseudococcidae Cockerell, 1905 ancestral family Phenacoleachiidae and is known in all primitive groups of mealybugs: Putoidae Tang, 1992: 382 (as a tribe of Pseudo- coccidae with several genera). Williams et g/g Puto, species-rich genera Heliococcus, al., 2011: 11 (as a separate monotypic family Coccura, Fonscolombia, Paraputo, Formi- without Ceroputo). Gavrilov-Zimin & Dan- cococcus and in many other genera, for ex- zig, 2012: 108 (synonymy). ample in legless mealybugs (g/g Antonina), Rhizoecidae Williams, 1969 (as a tribe of Pseu- species-rich genus Mirococcopsis, in numer- dococcidae). Hodgson, 2012: 8 (as a separate ous small and monotypic genera: Nudicau- family). Gavrilov-Zimin & Danzig, 2014: 124 da, Adelosoma, Antoninella, Cucullococcus, (synonymy). Lacombia, Ritsemia, Ehrhornia, Paratriony- Pennygullaniidae Koteja et Azar, 2008: 158. Gavrilov-Zimin & Danzig, 2012: 108 (syn- mus, Lankacoccus, Stricklandina, Madacan- onymy). thococcus, Mauricoccus, Mascarenococcus, Cirnecoccus, Pleistocerarius, etc. To date Two subfamilies of the Pseudococcidae more than 500 obligate ovoviviparous spe- generally differ in the following characters. cies of mealybugs from more than 60 genera Species of the subfamily Phenacoccinae have been found (Trapeznikova & Gavrilov, usually possess claw denticle, dorsal coni- 2008; I. Gavrilov-Zimin, unpublished), that cal setae, 9-segmented antennae, 5-locular is about 25% of the species diversity in the pores and only simple tubular ducts. Species global fauna. Moreover, there is no doubt of the subfamily Pseudococcinae without that the real number of ovoviviparous spe- claw denticles, with flagellate dorsal setae, © 2015 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 24(2): 236–259 244 I.A. GAVRILOV-ZIMIN. SYSTEM OF GENERIC GROUPS IN MEALYBUGS 6–8-segmented antennae and tubular ducts ability row of general taxonomic characters of different types (not only simple); 5-locu- with vast g/g Phenacoccus. lar pores absent. However, there are numer- Nominal genera (6): Eastia De Lotto, ous exclusions from this general definition. 1964 (Afrotropical), Mombasinia De Lotto, Moreover, some genera and groups of gen- 1964 (Afrotropical), Nairobia De Lotto, era (for example, g/g Metadenopus) have a 1964 (Afrotropical), Puto Signoret, 1876 questionable phylogenetic and taxonomic (mainly Holarctic), Rastrococcus Ferris, position – see the scheme of estimated re- 1954 (Oriental, Australasian), Trimerococ- lationships from Danzig & Gavrilov-Zimin, cus Balachowsky, 1952 (Palaearctic). 2014 with minor changes and additions Deviating genera/species. Several spe- (Fig. 4). cies of Puto (Palaearctic P. borealis Borch- senius, 1948 and Neotropical P. paramoensis System of the morphological Matile-Ferrero, 1985 + P. lamottei Matile- generic groups (g/g) Ferrero, 1985) and monotypic genera Mom- basinia and Nairobia lacked claw denticle. 1 g/g Puto Signoret, 1876 Moreover, Mombasinia lacked ostioles and demonstrates very peculiar large dorsal Combination of diagnostic characters groups of simple tubular ducts, enclosed by (plesiomorphic+apomorphic): claw with a multilocular pores. denticle; most or all cerarii with multiple All species of Rastrococcus have peculiar (more than 4) conical setae or all cerarii truncate conical setae. merged in a solid marginal band; 9-seg- mented antennae; both pairs of ostioles well 2 g/g Phenacoccus Cockerell, 1893 developed; complete anal apparatus (with one inner row of pores, one outer row of Combination of diagnostic characters spinulae and 6 long setae); tubular ducts of (plesiomorphic + apomorphic): cerarii simple type; trilocular pores numerous and each with 2–3 conical setae, never merged evenly scattered on all body surface. in marginal band; 9-segmented antennae; Additional characters: marginal cerarii quinquelocular pores present; both pairs of are usually supernumerary (more than 18 ostioles present. pairs) in comparison with other Phenacoc- Additional characters (plesiomorphic): cinae or even merged in the solid marginal well developed dorsal conical setae (often band of conical setae and trilocular pores organized in several medial cerarii), claw (as in Trimerococcus and in South African with a denticle; simple type of tubular ducts; endemic Eastia). well-developed legs and antennae, complete All species of the group are highly mov- anal apparatus; trilocular pores numerous able insects possessing well developed an- and evenly scattered on all body surface. tennae and legs and living openly on their Nominal genera (10): Coccura Šulc, 1908 host plants. Probably all species are ovovi- (Palaearctic), Fonscolombia Lichtenstein, viparous and do not construct wax ovisacs. 1877 (Palaearctic), Malekoccus Matile- There is no doubt that the group or at least Ferrero, 1988 (Palaearctic), Mammicoccus the genus Puto itself are the most ancient Balachowsky, 1959 (Neotropical), Octococ- and primitive mealybugs and neococcids cus Hall, 1939 (Afrotropical), Oxyacanthus (superfamily Coccoidea) as a whole. The De Lotto, 1970 (Afrotropical), Perystrix species of g/g Puto demonstrate all apomor- Gavrilov, 2004 (Palaearctic), Phenacoccus phic characters of the family Pseudococ- Cockerell, 1893 (global distribution), Stem- cidae, plesiomorphic characters inherited matomerinx Ferris, 1950 (Nearctic), Syna- from estimated ancestral family Phenaco- canthococcus Morrison, 1920 (Palaearctic leachiidae and almost uninterrupted vari- and Oriental). © 2015 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 24(2): 236–259 I.A. GAVRILOV-ZIMIN. SYSTEM OF GENERIC GROUPS IN MEALYBUGS 245 Usual complete anal apparatus Complicated anal apparatus Different examples of simplified anal apparatus Mirococcopsis ehrhornioidea Ritsemia pupifera Mirococcopsis kalaginae Cucullococcus arrabidensis Paratrionymus halocharis Ehrhornia fodiens Some peculiar types of anal apparatus Ripersiella brevipes Pararhodania armena Rhodania flava Fig. 3. Different types of anal apparatus. © 2015 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 24(2): 236–259