System of Generic Groups in Mealybugs (Homoptera: Coccinea: Pseudococcidae)

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 morphological 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.

Here I am providing a synopsis of the DISCUSSION elaborated system, a discussion of some additions to the system, basing on analysis of The main taxonomic characters used non-Palaearctic genera and a conspectus in the analysis of the genera (Fig. 1) 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-

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 C18 Vulva 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

Table 1. Wax glands in Pseudococcidae.

Discoidal wax glands

Simple discoidal pores Nudicauda nigra

Trilocular pores Cucullococcus Iberococcus

Quinquelocular pores

Multilocular pores

Pelionella Cylindrical wax glands

Simple tubular ducts

Coccidohystrix

Fungiform ducts

Oral collar ducts

Middle collar ducts

Collar tubular ducts

Double collar ducts

Crateriform ducts

Heliococcus

Mitral ducts 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 (С18) is reduced. 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

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- g/g Puto, species-rich genera Heliococcus, coccidae with several genera). Williams et 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. mus, Lankacoccus, Stricklandina, Madacan- Gavrilov-Zimin & Danzig, 2012: 108 (syn- thococcus, Mauricoccus, Mascarenococcus, onymy). 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,

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-segmented 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).

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.

Deviating genera/species. Majority of Nominal genera (6): Dawa Wil- deviating species are combined in the ge- liams, 1985 (Australasian), Peliococcus nus Fonscolombia, all species of which have Borchsenius, 1948 (global distribution), flagellate setae on dorsum, some species Peliococcop sis Borchsenius, 1948 (Palaearc- lack claw denticle or 5-locular pores or have tic), Erimococcus Ezzat, 1965 (Palaearctic), 6–8-segmented antennae. Monotypic genus Seyneria Goux, 1990 (Palaearctic), Pelio- Malekoccus, known from Saudi Arabia, dem- nella Kaydan, 2015 (Palaearctic). onstrates tubular ducts with flat ring near Deviating genera/species. The species of opening and peculiar large 5-locular pores, Seyneria have tubular ducts with collar. which are of the same size as multilocular pores. The species of Stemmatomerinx have 4 g/g Heliococcus Šulc, 1912 peculiar truncate conical setae with the base surrounded by small pores. The species of Combination of diagnostic characters Mammicoccus have unusual circuli, divided (unique apomorphic): crateriform tubular into protuberances. The correct taxonomic ducts. placing of the genus Octococcus (4 species) Additional characters (plesiomorphic): is most disputable: all species have only 2 claw with a denticle; cerarii each with 2–3 (posterior) pairs of cerarii, dorsal flagellate conical setae; 9-segmented antennae; quin- setae and ducts with flat ring (rim) near quelocular pores; dorsal surface of body opening; two species (including the type covered by conical setae; both pairs of osti- one) have ducts with two rims (additional oles present; trilocular pores numerous and rim encircling the middle of the duct). evenly scattered on all body surface. The g/g Heliococcus includes only one 3 g/g Peliococcus Borchsenius, 1948 genus in the global fauna. This genus is probably a collateral blind branch of the phylo- Combination of diagnostic characters genetic line Puto-Phenacoccus (see Fig. 4). (apomorphic): clusters of multilocular The species of Heliococcus show in addition pores and/or tubular ducts and/or with en- to their general plesiomorphic morphology larged conical setae with attached trilocu- and a plesiomorphic mode of life (highly lar pore(s) just near the base. Both these movable ovoviviparous females) one unique characters are present in the type species apomorphic character – the appearance of of Peliococcus, P. chersonensis (Kiritshenko, crateriform tubular ducts. 1936), and also in several other species of Deviating genera/species: some species Peliococcus and related genera. Most other (for example, H. takae (Kuwana, 1907), species of g/g Peliococcus show only one of H. takahashii (Kanda, 1935) and H. vari- these characters: either clusters of glands oporus Matesova, 1968) have cerarii with or setae with pore just near the base that multiple conical setae. impedes a classification of the group and a practical identification of the taxa. 5 g/g Coccidohystrix Lindinger, 1943 Additional characters (plesiomorphic): well developed dorsal conical setae (often Combination of diagnostic charac- organized in several medial cerarii), simple ters (apomorphic): cerarii replaced by the type of tubular ducts, well developed legs groups of large conical setae without as- and antennae, complete anal apparatus; sociated trilocular or quinquelocular pores claw with a denticle; cerarii each with 2–3 (mamelons); tubular ducts of characteristic conical setae, never merged in marginal bottle-shaped form; the presence of more band; 9-segmented antennae; quinquelocu- or less cylindrical setae on dorsum; the ab- lar pores; both pairs of ostioles present; sence of ostioles. trilocular pores numerous and evenly scat- Additional characters (plesiomorphic): tered on all body surface. claw with a denticle; 9-segmented antennae;

© 2015 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 24(2): 236–259 I.A. GAVRILOV-ZIMIN. SYSTEM OF GENERIC GROUPS IN MEALYBUGS 247 quinquelocular pores present ; tendency to dorsal flagellate setae. Monotypic genus decreasing of the number of trilocular pores. Paramococcus lacks both pairs of ostioles. Nominal genera (3): Coccidohystrix Some species of Heterococcus lack cerarii Lindinger, 1943 (Palaearctic + Oriental), and have dorsal flagellate setae. Breven- Lantanacoccus Williams et Granara de Wil- nia dasiforae (Danzig, 1977) and B. rehi link, 1992 (Neotropical), Eriocorys De Lot- (Lindinger, 1943) lack claw denticle. to, 1967 (Afrotropical). Deviating genera/species. Some of Coc- 7 g/g Boreococcus Danzig, 1960 cidohystrix species demonstrate significant Combination of diagnostic characters decreasing of the number of trilocular pores, (unique apomorphic): trilocular pores lack- up to their presence near spiracles only. This ed and replaced by multilocular pores of feature in combination with the absence of two types: usual type and asteroid multi- ostioles and scattering of enlarged dorsal locular pores. conical setae denote similarity and prob- Additional characters (plesiomorphic): ably relationship of Coccidohystrix g/g with claw with a denticle; cerarii each with 2–3 the family Eriococcidae (felt scales) and conical setae; 9-segmented antennae; both it seems that Coccidohystrix can be con- pairs of ostioles present. sidered as an ancestor group for the felt Nominal genera (2): Annulococcus Ja- scales. In particular, two recently described mes, 1936 (Afrotropical), Boreococcus Dan- North-African species, C. maghribiensis and zig, 1960 (Palaearctic). C. monicae (Gavrilov-Zimin & Matile-Fer- Deviating genera/species. Both species rero, 2014) differ, in fact, from the species of Annulococcus have collar tubular ducts of the largest (and probably most primitive) and numerous long flagellate setae on dor- genus of felt scales, Acanthococcus Signoret, sum. 1875, only in the presence of small number of trilocular pores near spiracles. 8 g/g Mirococcus Borchsenius, 1947 Lantanacoccus lacked claw denticle and has 6-segmented antennae. Combination of diagnostic characters Coccidohystrix insolita (Green, 1908) (apomorphic): cerarii totally lacked. and monotypic genus Eriocorys save one Additional characters (plesiomorphic + (posterior) pair of ostioles. apomorphic): claw with a denticle; quinquelocular pores usually present; anal appa- 6 g/g Heterococcus Ferris, 1918 ratus often simplified; tendency to decreasing of the number of trilocular pores. Combination of diagnostic charac- I include in this group different mealy- ters (apomorphic): trilocular pores totally bugs with features of simplification and re- lacked or very few and present near spiracles duction of main morphological characters, and in the cerarii only; quinquelocular pores such as cerarii, ostioles, anal apparatus, wax numerous and scattered on all body surface. glands, etc. These mealybugs are placed in Additional characters (plesiomorphic): different small and monotypic genera and claw with a denticle; cerarii each with 2–3 perhaps form not monophyletic group. This conical setae; dorsal setae usually conical; group is probably originated from different 9-segmented antennae. ancestors similar with modern species of Nominal genera (4): Brevennia Goux, Fonscolombia or/and Brevennia (subfam- 1940, Heterococcus Ferris, 1918, Paramococ- ily Phenacoccinae), that confirmed by the cus Foldi et Cox, 1989 (Neotropical), Lain- presence of claw denticle, often presence giococcus Morrison, 1945 (Australasian), of quinquelocular pores, total or partial re- Deviating genera/species. Some species placement of trilocular pores by quinque- of Brevennia lack anterior ostioles and have locular ones, 8–9 (rarely 7)-segmented an-

© 2015 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 24(2): 236–259 248 I.A. GAVRILOV-ZIMIN. SYSTEM OF GENERIC GROUPS IN MEALYBUGS tennae and simple tubular ducts. The genus nae segments, appearing of collar tubular Mirococcus seems to be the most primitive in ducts and flagellate setae on dorsum. considered group, because it saves usual for A row of nominal monotypic and oligo- most mealybugs high number of evenly dis- typic genera, described mainly from south- tributed trilocular pores, more or less com- ern hemisphere do not have any clear dif- plete anal apparatus and thick setae, accom- ferences from the oldest genera Paraputo or panied by a group of trilocular pores in place Formicococcus and the names of such genera of last (18-th) pair of cerarii (pseudocerarii). must be probably synonymised in future; Nominal genera (8): Cucullococcus Fer- this supposed synonymy is noting below. ris, 1941 (Holarctic), Discococcus Ferris, Nominal genera (29): Anisococcus Fer- 1953 (Nearctic and Neotropical), Ehrhorn- ris, 1950 (Nearctic, Neotropical), Brasiliputo ia Ferris, 1918 (Holarctic), Mollicoccus Wil- Williams et Granara de Willink, 1992 (Neo- liams, 1960 (Australasian), Neosimmondsia tropical, ?=Paraputo), Chileputo Williams Laing, 1930 (Australasian), Scaptococcus et Granara de Willink, 1992 (Neotropical, McKenzie, 1964 (Nearctic), Sinococcus Wu ?=Paraputo), Crenicoccus Williams, 2004 et Zheng, 2000 (Palaearctic), Stipacoccus (Oriental, ?=Formicococcus), Criniticoccus Tang, 1992 (Palaearctic). Williams, 1960 (Australasian), Cyperia De Deviating genera/species. Three oligo- Lotto, 1964 (Afrotropical), Delococcus Fer- typic genera, Cucullococcus, Discococcus, ris, 1955 (Afrotropical), Eucalyptococcus Ritsemia and Sinococcus are characterized Williams, 1985 (Australasian, ?=Paraputo), by very peculiar morphological characters Exallomochlus Williams, 2004 (Oriental), and perhaps related to each other. All these Exilipedronia Williams, 1960 (Oriental), genera demonstrate sclerotization of the Farinococcus Morrisson, 1922 (Neotropical, cuticle on whole body or on some parts of ?=Paraputo), Formicococcus Takahashi, 1928 body in adult females, significant reduction (Palaearctic, Oriental), Hopefoldia Foldi, of the number of trilocular pores, absence 1988 (Neotropical), Lachnodiella Hem- of ostioles and presence of numerous circuli pel, 1910 (Neotropical, ?=Formicococcus), (4–6 in number). All species of Discococcus Laminicoccus Williams, 1960 (Australasian, lack claw denticle. ?=Paraputo), Lanceacoccus Williams, 2004 (Oriental), Lankacoccus Williams, 2004 9 g/g Paraputo Laing, 1929 (Oriental), Mascarenococcus Mamet, 1940 (Afrotropical), Moystonia Williams, 1985 Combination of diagnostic characters (Australasian, Oriental), Mutabilicoccus (plesiomorphic + apomorphic): cerarii Williams, 1960 (Australasian), Odacoccus with multiple conical setae, which are often Williams et Watson, 1988 (Australasian), merged in entire marginal band; dorsal sur- Paraputo Laing, 1929 (Palaearctic, Orien- face of body usually covered by conical se- tal), Circaputo McKenzie, 1962 (Nearctic), tae, more rarely – by thick flagellate setae; Palmicultor Williams, 1960 (pantropical), quinquelocular pores absent, claw without Pleistocerarius Matile-Ferrero, 1970 (Af- a denticle. rotropical), Stricklandina Matile-Ferrero, Additional characters (plesiomorphic + 1998 (Afrotropical, Oriental), Sarococcus apomorphic): tubular ducts with collar; both Williams et de Boer, 1973 (Australasian), pairs of ostioles well developed; 6–8-seg- Tomentocera Beardsley, 1964 (Australasian: mented antennae; trilocular pores numerous Hawaii), Yudnapinna Williams, 1985 (Aus- and evenly scattered on all body surface. tralasian). The group is undoubtedly most primi- Deviating genera/species. The species tive in subfamily Pseudococcinae and shares of Formicococcus and Lachnodiella have many characters with Puto, but lacked claw numerous (more than 6–8) setae of anal denticle, decreasing of the number of anten- ring. Monotypic Lankacoccus demonstrates

9-segmented antennae and numerous 5-lo- 11 g/g Nipaecoccus Šulc, 1945 cular pores on venter. The species of Moys- tonia and Tomentocera have small fungiform Combination of diagnostic characters ducts. Monotypic genus Circaputo demon- (apomorphic): conical setae (in addition or strates minute claw denticle. instead of cerarian setae) numerous along all Correct taxonomic placing of monotypic body margin or/and throughout on dorsum. Hopefoldia is very questionable; this genus Additional characters (plesiomorphic): is characterized by very unusual sclerotized claw without a denticle; quinquelocular plates with pores, so-called “poraria”, scat- pores absent; antennae 6–8-segmented; cer- tered on both body sides. arii (if present) each with 2–3 conical setae; trilocular pores numerous and evenly scattered on all body surface. 10 g/g Pedronia Green, 1922 The g/g Nipaecoccus is similar with g/g Combination of diagnostic characters Paraputo in the presence of numerous dor- (apomorphic): dorsal enlarged conical setae sal conical setae in addition to conical setae located on sclerotized tubercles and/or on of cerarii. prominent lobes. The type species of the genera Hypogeo- Additional characters (plesiomorphic): coccus and Nipaecoccus, namely H. barbarae claw without a denticle; quinquelocular Rau, 1938 and N. nipae (Maskell, 1893) are pores absent; antennae 6–8-segmented. very similar morphologically and the char- Nominal genera (9): Acrochordonus Cox, acters of other nominal species of both gen- 1987 (Australasian: New Zealand), Agas- era are totally overlapped. tococcus Cox, 1987 (Australasian: New In the review of Palaearctic fauna (Dan- Zealand), Clavicoccus Ferris, 1948 (Aus- zig & Gavrilov-Zimin, 2014, 2015) we con- tralasian: Hawaii), Extanticoccus Williams, sidered the genera Adelosoma and Trabutina 2004 (Oriental), Neoclavicoccus Cohic, in the separate g/g Trabutina, but basing on 1959 (Australasian: New Caledonia), Ne- the present analysis of the world fauna we sopedronia Beardsley, 1971 (Australasian: do not see morphological hiatus between it Hawaii), Pedrococcus Mamet, 1942 (Afro- and g/g Nipaecoccus. tropical, Oriental), Pedronia Green, 1922 Nominal genera (22): Adelosoma Borch- (Oriental), Tylococcus Newstead, 1897 (Af- senius, 1948 (Palaearctic), Amonostherium rotropical, Oriental). Morrison et Morrison, 1922 (Nearctic), Aus- Deviating genera/species. Two endemic tralicoccus Williams, 1985 (Australasian), New Zealand genera, Acrochordonus are Conulicoccus Williams, 1985 (Australasian), Agastococcus as well as Afrotropical Tylo- Cyphonococcus Cox, 1987 (Australasian), coccus are characterized by peculiar very Erioides Green, 1922 (Oriental), Epicoccus prominent lateral lobes on posterior ab- Cockerell, 1982 (Australasian), Fijicoccus dominal segments or along all body mar- Williams et Watson, 1988 (Australasian), gin; these lobes bear multiple conical se- Grewiacoccus Brain, 1918 (Afrotropical), tae; similar, but smaller lobes are present Hordeolicoccus Williams, 2004 (Orien- in Oriental monotypic Extanticoccus. Type tal), Hypogeococcus Rau, 1938 (Nearctic, species of Hawaiian endemic genus Clavi- Neotropical, Afrotropical, Australasian), coccus also have numerous extremely long Kenmorea Williams, 1985 (Australasian), dorsal prominences with multiple conical Lanceacoccus Williams, 2004 (Oriental), setae; the second species of the genus have Madangiacoccus Williams, 1985 (Austral- prominences of moderate size, but also with asian), Melanococcus Williams, 1985 (Aus- multiple conical setae. Probably, these gen- tralasian), Nipaecoccus Šulc, 1945 (global era occupy intermediate position between distribution), Paramonostherium Williams, g/g Paraputo and g/g Pedronia. 1985 (Australasian), Porococcus Cockerell,

1898 (Neotropical), Pseudoripersia Cock- g/g Pseudococcus). In view of these facts erell, 1899 (Australasian), Strombococcus we consider all g/g Planococcus, genera in- Williams, 1985 (Australasian), Trabutina side of it and also the tribe Planococcini Ez- Marchal, 1904 (Palaearctic), Ventrispina zat et McConnell, 1956 as unnatural. Williams, 1985 (Australasian). Nominal genera (45): Anthelococcus Deviating genera/species. McKenzie, 1964 (Nearctic), Asaphococ- The taxonomic position of the genus cus Cox, 1987 (Australasian), Asteliacoc- Amonostherium is disputable; its type spe- cus Williams, 1985 (Australasian), Bonino- cies has a claw denticle and lack ostioles. coccus Kawai, 1973 (Australasian: Bonin The species of Cyphonococcus have numer- Islands, ?=Trionymus), Callitricoccus Wil- ous fungiform ducts. liams, 1985 (Australasian), Chaetotriony- mus Williams, 1985 (Australasian, ?= Tri- 12 g/g Trionymus Berg, 1899 onymus), Chlorococcus Beardsley, 1971 (Australasian: Hawaii), Chnaurococcus Fer- Combination of diagnostic characters ris, 1950 (Nearctic, ?=Trionymus), Cirne- (apomorphic): cerarii with 2–3 conical se- coccus Mamet, 1967 (Afrotropical), Crisi- tae; conical setae present in cerarii only; coccus Ferris, 1950 (global distribution), both body sides covered by flagellate or Crocydococcus Cox, 1987 (Australasian), hair-like setae. Delottococcus Cox et Ben-Dov, 1986 (Afro- Additional characters (plesiomorphic): tropical), Dysmicoccus Ferris, 1950 (global claw without a denticle; antennae 6–8-seg- distribution), Erium Cockerell, 1897 (Aus- mented; trilocular pores numerous and tralasian), Eurycoccus Ferris, 1950 (global evenly scattered on all body surface. distribution, ?=Trionymus), Ferrisia Fulla- A row of nominal monotypic and oligo- way, 1923 (global distribution), Ityococcus typic genera, described mainly from south- Williams, 1985 (Australasian), Leptococcus ern hemisphere do not have any clear dif- Reyne, 1961 (Oriental), Macrocepicoccus ferences from the older genus Trionymus and Morrison, 1919 (Neotropical), Maculicoc- the names of such genera must be probably cus Williams, 1960 (Australasian), Mada- synonymised in future; this supposed syn- gasia Mamet,1962 (Afrotropical), Mar- onymy is noting below. endellea De Lotto, 1967 (Afrotropical), In the review of Palaearctic fauna Mauricoccus Mamet, 1967 (Afrotropical), (Danzig & Gavrilov-Zimin, 2014, 2015) Nesococcus Ehrhorn, 1916 (Australasian), we considered the genera Planococcus and Ohiacoccus Beardsley, 1971 (Australasian), Crisicoccus in the separate g/g Planococcus. Oracella Ferris, 1950 (Nearctic), Orococcus This group included species which are very De Lotto, 1964 (Afrotropical), Orstomi- similar morphologically with the species coccus Mamet, 1962 (Afrotropical), Para- from large genera Trionymus Berg, 1899 and doxococcus McKenzie, 1962 (Nearctic), Dysmicoccus Ferris, 1950, but demonstrate Paraferrisia Williams et de Boyer, 1973 so-called “anal bar” – sclerotized stripe on (Australasian), Phyllococcus Ehrhorn, 1916 ventral surface of anal lobe. Such character (Australasian: Hawaii), Planococcus Ferris, is very subjective, because it is often diffi- 1950 (global distribution), Poecilococcus cult to differ anal bar from other variants Brookes, 1981 (Australasian), Pseudofer- of cuticular sclerotization on anal lobes. risia Kaydan et Gullan, 2012 (Nearctic), Also, examples of individual variation of Pseudotrionymus Beardsley, 1971 (Austral- the character, i.e. presence/absence of the asian: Hawaii), Renicaula Cox, 1987 (Aus- anal bar in females of the same population, tralasian), Spartinacoccus Kosztarab, 1996 are known, for example, in the type series (Nearctic), Strandanna De Lotto, 1969 of Atrococcus expressus (Borchsenius, 1949) (Afrotropical), Tasmanicoccus Williams, and A. pacificus (Borchsenius, 1949) (see 1985 (Australasian), Telocorys De Lotto,

1967 (Afrotropical), Trionymus Berg, 1899 Trionymus!) and has heavily sclerotized lo- (global distribution), Trochiscococcus Wil- bate posterior end of body (two posterior liams et Pellizzari, 1997 (Afrotropical), Tur- segments); the other abdominal segments binococcus Beardsley, 1966 (Australasian), significantly narrowed in comparison with Tympanococcus Williams, 1967 (Oriental), cephalothorax. Villosicoccus Williams, 1985 (Australasian). Deviating genera/species. Monotypic 13 g/g Pseudococcus Westwood, 1840 Anthelococcus has dorsal conical setae (together with flagellate ones) and 5-locular Combination of diagnostic characters pores on venter. Four monotypic and oligo- (apomorphic): fungiform ducts typic genera, Maculicoccus, Pseudotriony- Additional characters (plesiomorphic): mus, Spartinacoccus, Trochiscococcus save claw without a denticle; cerarii with 2–3 only several cerarian pairs, but these cerarii conical setae; antennae 6–8-segmented; with multiple conical setae, the character trilocular pores numerous and evenly scat- which these genera share with g/g Paraputo. tered on all body surface. Monotypic Cirnecoccus demonstrates Nominal genera (13): Apodastococcus unusually prominent and sclerotized anal Williams, 1985 (Australasian), Atrococ- lobes and replacing of most cerarii (exclud- cus Goux, 1941 (Holarctic), Atriplicicoc- ing only C18) by long flagellate setae. Simi- cus Williams et Granara de Willink, 1992 lar in size, but not sclerotized anal lobes is (Neotropical), Chorizococcus McKenzie, known only in monotypic Poecilococcus. 1960 (Nearctic), Chryseococcus Cox, 1987 Monotypic Marendellea has peculiar (Australasian), Colombiacoccus Williams clusters of multilocular pores+tubulars et Granara de Willink, 1992 (Neotropical), ducts, the same as in g/g Peliococcus, but Cormiococcus Williams, 1989 (Afrotropi- share most of other characters with the ge- cal), Distichlicoccus Ferris, 1950 (Nearctic, nus Trionymus. Neotropical), Gallulacoccus Beardsley, 1971 Ferrisia and very similar monotypic ge- (Australasian), Maconellicoccus Ezzat, 1958 nus Pseudoferrisia, have unique tubular (pantropical), Papuacoccus Williams et ducts with sclerotized rim, bearing several Watson, 1988 (Australasian), Paracoccus pores and flagellate setae. Monotypic Para- Ezzat et McConell, 1956 (pantropical), Spi- ferrisia has tubular ducts with similar scler- lococcus Ferris, 1950 (Nearctic). otized rim, but without pores or setae on it. Deviating genera/species. The species Monotypic Londiania has 6 pairs of with of Chorizococcus have claw denticle. The multiple very thin (almost flagellate) setae. species of Maconellicoccus possess 9-seg- Monotypic Madagasia and Telocorys mented antennae. Monotypic Cormiococcus possess 17–19 pairs of pseudocerarii, each has very numerous multilocular pores, scat- with 2–3 long flagellate setae. tered on all body surface and even inside of The species of Renicaula have very small spiracular peritremae. antennae and legs and enlarged hind coxae with numerous translucent pores. 14 g/g Neotrionymus Borchsenius, 1948 The species of Chlorococcus Beardsley, 1971 (Australasian: Hawaii), Leptococcus Combination of diagnostic characters Reyne, 1961 (Oriental), Macrocepicoccus (apomorphic): total or partial replacement have unusually long, slender antennae and of trilocular pores by multilocular ones in legs with long pointed claws. combination with a presence of one several The taxonomic position of monotypic normally developed cerarian pairs. genus Phyllococcus, endemic of Hawaii, is Additional characters (plesiomorphic): questionable; the type species produces claw without a denticle; cerarii with 2–3 peculiar galls (very rare example in g/g conical setae; antennae 6–8-segmented.

Iberococcus Rhizoecus Nudicauda ? group of genera 54 53 Allomirmococcus ? Mirococcopsis 55 group of genera (?=Humococcus) 52 Cryptoripersia 51 Rhodania

35 Archanginella Metadenopus Kalaginella 50 Antonina Balanococcus 35 group of genera Cucullococcus Paratrionymus 46 30 45 Miscanthicoccus Kiritshenkella 29 Ritsemia 35 ? 48 47 Neotrionymus Trionymus Spilococcus 28 49 (?=Eurycoccus) Sinococcus Chorizococcus 44 Ehrhornia Misericoccus 27 Planococcus 42 43 Atrococcus Discococcus 40 26 ? 42 Heterococcus Dysmicoccus Pseudococcus 41 (?= Paracoccus) 25 Pararhodania 22 Stipacoccus ? 20 Brevennia 37 24 Pedronia 36 Mirococcus 36a Clavicoccus Pedrococcus Formicococcus 21 Boreococcus Tylococcus Lachnodiella 39 34 38 32 23 20 Paraputo Nipaecoccus 37 33 group of genera Lankacoccus Lanceacoccus Fonscolombia Eriococcidae 19 Acanthococcus Perystrix

11 18 Seyneria Coccura Synacanthococcus Erimococcus Pelionella 17 13 9 15 10 8 12 16 Coccidohystrix Phenacoccus Peliococcus 10a Lantanacoccus 14 Peliococcopsis

Trimerococcus Rastrococcus Eastia 4 5 Heliococcus 7 6 31 (CeroputoPuto=Australiputo)

3 extinct ancestor Phenacoleachiidae

2 1 Xenococcidae ? Margarodidae s.l.

Nominal genera (6): Coorongia Wil- 15 g/g Metadenopus Šulc, 1933 liams, 1985 (Australasian), Miscanthicoc- cus Takahashi, 1957 (Holarctic), Neoriper- Combination of diagnostic characters sia Kanda, 1943 (Palaearctic, Australasian: (apomorphic): total replacement of triloc- Bonin Islands), Neotrionymus Borchsenius, ular pores by multilocular/quinquelocular 1948 (Palaearctic), Pandanicola Beardsley, ones in combination with the absence of 1966 (Australasia), Rosebankia De Lotto, cerarii. 1967 (Afrotropical). Additional characters (plesiomorphic): Deviating genera/species. Monotypic claw without a denticle; antennae 6-8-seg- Miscanthicoccus possesses peculiar campan- mented. iform spiracles. Both Miscanthicoccus and According to these characters the group Neoripersia have zones of shagreen cuticle. occupies intermediate position between g/g Monotypic Rosebankia demonstrates multi- Neotrionymus and g/g Mirococcopsis. Among locular pores of two sizes on both body sides. 8 genera considered below, five,Cypericoc-

Fig. 4. Scheme of estimated relationships between main generic groups of Pseudococcidae and related families. Ovals designate borders of families and generic groups. Species-rich and/or phylogentically important genera are written in larger font.

Evolutional changes of characters: 1 – lack of compound discoidal wax glands, primarily absence of ostioles and tubular ducts, primarily presence of female pupal larva; 2 – appearance of ostioles and swirled trilocular pores; 3 – lack of abdominal spiracles, appearance of cerarii and tubular ducts; 4 – truncate conical setae; 5 – solid marginal band of conical setae and trilocular pores; 6 – standardization of the number of cerarian setae (up to 2 per cerarius) and of the size of trilocular pores (equal on all body); 7 – crateriform ducts; 8 – enlarging of conical setae; 9 – tubular ducts with associated simple pores just near the excretory opening;10 – lacking of ostioles and normal cerarii, appearance of mamelons and “bottle-shaped” tubular ducts, decreasing of the number of trilocular pores; 10а – lacking of claw denticle; 11 – total lacking of trilocular pores; 12 – clusters of multilocular pores and/or tubular ducts;13 – tubular ducts with collar;14 – dorsal 5-locular pores;15 – dorsal clusters include tubular ducts only;16 – multilocular pores of 2 different types; 17 – tubular ducts forming marginal band and produce acetabuliform ovisac;18 – decreasing of the number of cerarii and appearance of dorsal flagellate setae; 19 – pseudocerarii along body margin;20 – decreasing of the number of 3-locular pores and their partial replacement by other discoidal glands; 21 – trilocular pores replaced by multilocular pores, tubular ducts with collar; 22 – total lacking of trilocular pores; 23 – total lacking of cerarii and simplification of anal apparatus; 24 – dorsal setae conical, lacking 5-locular pores; 25 – enlargement of anal ring and appearance of unusual pores on it; 26 – multiple circuli with double fringe, cuticle sclerotized; 27 – mitral ducts, lacking of 5-locular pores; 28 – conical setae on dorsum; 29 – lacking of tubular ducts; 30 – reduction of legs, trilocular pores without fringe; 31 – lacking of claw denticle; 32 – decreasing of the number of cerarii; 33 – enlarging of dorsal conical setae; 34 – dorsal conical setae located on sclerotized tubercles; 35 – lacking of cerarii; 36 – decreasing of the number of 3-locular pores; 36a – prominent lateral lobes of body; 37 – decreasing and standardization of the number of conical setae in cerarii; 38 – anal apparatus with numerous (more than 8) setae; 39 – tubular ducts of simple type only; 40 – anal lobe bar present; 41 – fungiform ducts; 42 – lacking of auxiliary flagellate setae in cerarii and decreasing of the number of cerarii; 43 – secondary appearance of claw denticle; 44 – fungiform ducts of different sizes; 45 – grouping of 3-locular pores in medial zone of body; 46 – multilocular pores form non-interrupted marginal band; 47 – replacement of trilocular pores by multilocular pore; 48 – campaniform spiracles; 49 – conical setae on dorsum; 50 – reducing of legs and sclerotization of cuticle; 51 – complete anal apparatus; 52 – simplification of anal apparatus; 53 – tubular duct opening encircled by a flat sclerotized ring; 54 – simplification of anal apparatus together with enlarging of scanty pores and spinulae, appearance of bitubular and tritubular ducts; 55 – enlargement of ostioles, legs and antennae, lacking of tubular ducts.

© 2015 Zoological institute, Russian Academy of Sciences, Zoosystemitica Rossica 24(2): 236–259. 254 I.A. GAVRILOV-ZIMIN. SYSTEM OF GENERIC GROUPS IN MEALYBUGS cus, Metadenopus, Misericoccus, Paratriony- Borchsenius, 1962 (Palaearctic), Cryptorip- mus and Syrmococcus show replacement ersia Cockerell, 1899 (Nearctic, Afrotropi- of trilocular pores by multilocular ones. cal, Australasian), Diversicrus De Lotto, Two else genera, Rhodania and monotypic 1970 (Afrotropical), Glycycnyza Danzig, Archanginella, lack multilocular pores as 1974 (Palaearctic), Hadrococcus Williams, well as trilocular ones, but have numerous 1985 (Australasian), Humococcus Ferris, quinquelocular pores on both body sides. 1953 (Neactic, Neotropical, Australasian), The fifth genus, monotypic Kalaginella, has Iberococcus Gómez-Menor Ortega, 1928 only quinquelocular pores, which similar in (Palaearctic), Inopicoccus Danzig, 1971 (Pa- size with multilocular pores of other mealy- laearctic), Lacombia Goux, 1940 (Palaearc- bugs. Furthermore, large discoidal pores of tic), Lenania De Lotto, 1964 (Afrotropical), complicated structure are demonstrated by Madeurycoccus Mamet, 1959 (Afrotropical), Rhodania aeluropi Williams et Moghaddam, Mirococcopsis Borchsenius, 1948 (Palaearc- 2007 which differs in this and in some other tic), Mizococcus Takahashi, 1928 (Oriental), characters from its cogeneres and from all Natalensia Brain, 1915 (Afrotropical), Nu- other species of g/g Metadenopus. dicauda Gavrilov, 2006 (Palaearctic), Palau- The name of American genus Misericoc- coccus Beardsley, 1966 (Australasian), Para- cus is probably junior synonym of Paratri- discococcus Williams, 1985 (Australasian), onymus. Pilococcus Takahashi, 1928 (Oriental), Nominal genera (8): Archanginella Dan- Prorsococcus Williams, 1985 (Australasian), zig et Gavrilov-Zimin, 2013 (Palaearctic), Tridiscus Ferris, 1950 (USA), Volvicoccus Cypericoccus Williams, 1985 (Australasian), Goux, 1945 (Palaearctic). Kalaginella Danzig et Gavrilov-Zimin, 2013 Deviating genera/species. Monotypic (Palaearctic), Metadenopus Šulc, 1933 (Pa- Diversicrus possesses unusual enlarged hind laearctic), Misericoccus Ferris, 1953 (Nearc- legs. tic), Paratrionymus Borchsenius, 1948 (Pa- laearctic), Rhodania Goux, 1935 (Palaearc- 17 g/g Rhizoecus Künkel d’Herculais, 1878 tic), Syrmococcus Ferris, 1953 (Nearctic). Deviating genera/species. The type spe- Combination of diagnostic characters cies of both Misericoccus and Paratrionymus (apomorphic): special type of anal appara- have occasional conical setae on last ab- tus with enlarged pores and spinulae; bases dominal tergite. of antennae located closely to each other; cephalic sclerotized plate and bi-/tritubular 16 g/g Mirococcopsis Borchsenius, 1948 ducts usually present; antennae 5–6-segmented. Combination of diagnostic characters Additional characters (plesiomorphic): (apomorphic): cerarii and any conical setae claw without a denticle; trilocular pores nu- absent. merous and evenly distributed; anal appara- Additional characters (plesiomorphic): tus often simplified; cerarii absent (with the claw without a denticle; legs normally de- only exclusion in Prorhizoecus). veloped; antennae 6–8-segmented; trilocu- The g/g Rhizoecus is one of the most spe- lar pores numerous and evenly distributed; cialized and aberrant groups in the family anal apparatus often simplified. Pseudococcidae, but the group shares with Nominal genera (24): Acaciacoccus Wil- other mealybugs obvious unique synapo- liams et Matile-Ferrero, 1994 (Afrotropical), morphic characters of Pseudococcidae: pres- Antoninella Kiritshenko, 1937 (Palaearctic), ence of ostioles and trilocular pores and also Gomezmenoricoccus Kozár et Walter, 1985 non-unique synapomorphies: similar struc- (Palaearctic), Bimillenia Matile-Ferrero et ture of tubular ducts and multilocular pores Ben-Dov, 1999 (Palaearctic), Coleococcus and similar structure of anal apparatus.

Nominal genera (15): Benedictycoccina several exclusions); anal apparatus often Kozár et Foldi, 2004 (Neotropical, Afro- simplified; cerarii usually absent. tropical), Brevicoccus Hambleton, 1946 The world fauna of mealybugs includes (Neotropical), Capitisetella Hambleton, at least 26 genera, which is characterized 1977 (Neotropical), Coccidella Hamble- by strong reduction or loss of legs. Eight of ton, 1946 (Neotropical), Geococcus Green, these genera are considered by most modern 1902 (global distribution), Hambleton- specialists to be closely related and placed rhizoecus Kozár et Konczné Benedicty, in separate tribe Serrolecaniini Shinji, 1935. 2005 (Neotropical), Kissrhizoecus Kozár The diagnostic characters of this tribe are et Konczné Benedicty, 2005 (Palaearctic), caudally directed vulva and groups of mi- Leptorhizoecus Williams, 1998 (Oriental), crotubular ducts located on ventral cuticle Marottarhizoecus Kozár et Konczné Bene- in place of reduced legs or on the surface of dicty, 2002 (Afrotropical), Pseudorhizoecus peculiar bag-like structures (probably mod- Green, 1933 (Neotropical), Pygmaeococcus ified hind coxae). The tribe includes 8 gen- McKenzie, 1960 (USA), Prorhizoecus Mill- era, seven of which are mainly Palaearctic: er et McKenzie, 1971 (Nearctic), Rhizoecus Paraserrolecanium, Idiococcus, Porisaccus, Kunckel d’Herculais 1878 (global distri- Paraporisaccus, Serrolecanium, Chaetococ- bution), Ripersiella Tinsley, 1899 (global cus, Tangicoccus and one monotypic genus distribution), Williamsrhizoecus Kozár et Kermicus has Oriental areal. Other genera Konczné Benedicty, 2007 (Neotropical, Af- of “legless” mealybugs perhaps not closely rotropical). related to each other and to Serrolecaniini Deviating genera/species. The species of and demonstrate convergent similarity only Geococcus have prominent sclerotized anal (see more detail discussion in Hendricks & lobes, each with stout conical seta at apex. Kosztarab, 1999). Monotypic Kissrhizoecus has numerous Nominal genera (26): Albertinia De quinquelocular pores on both body sides Lotto, 1970 (Afrotropical), Aemulantonina (unique example in g/g Rhizoecus). Mono- Williams, 2004 (Oriental), Acinicoccus Wil- typic Pseudorhizoecus possesses simplified liams, 1985 (Australasian), Antonina Signo- anal apparatus, without any pores and spi- ret, 1872 (global distribution), Antoninoi- nulae. des Ferris, 1953 (Nearctic), Chaetococcus Monotypic Prorhizoecus is probably Maskell, 1893 (mainly Nearctic), Chloeoon most primitive member of the group; it Anderson, 1788 (Oriental), Conicosoma De saves usual type anal apparatus (with small Lotto, 1970 (Afrotropical), Cypericoccus pores and spinular) and normally developed Williams, 1985 (Australasian), Idiococcus Takahashi et Kanda, 1939 (Palaearctic), C18, but has antennae similar with that in other species of the group and peculiar bul- Kermicus Newstead, 1897 (Oriental), Mi- bous ducts like in some other genera of the conicoccus Williams et Miller, 1999 (Neo- group, for example, in Pygmaeococcus. tropical), Nesticoccus Tang, 1977 (Palaearc- tic), Paludicoccus Ferris, 1918 (Nearctic), 18 g/g Antonina Signoret, 1872 Parapaludicoccus Mamet, 1962 (Afrotropi- cal), Paraporisaccus Lu et Wu, 2011 (Pa- Combination of diagnostic characters laearctic), Paraserrolecanium Wu, 2010 (apomorphic): legs strongly reduced or ab- (Palaearctic), Paulianodes Mamet, 1953 sent at all; hind coxae enlarged, often modi- (Afrotropical), Peridiococcus Williams, fied to sclerotized bag-like structures, coved 1985 (Australasian), Porisaccus Hendricks by microtubular ducts or translucent pores. et Kosztarab, 1999 (Palaearctic), Pseudan- Additional characters (plesiomorphic): tonina Green, 1922 (Oriental, Nearctic, claw without a denticle; trilocular pores nu- Neotropical), Serrolecanium Shinji, 1935 merous and evenly distributed (with only (Palaearctic), Sphaerococcus Maskell, 1892

(Australasian, Afrotropical), Tangicoc- Agastococcus – g/g Pedronia cus Kozár et Walter, 1985 (Palaearctic), Albertinia – g/g Antonina Quadrigallicoccus Williams et Miller, 1999 Allomyrmococcus – g/g Allomyrmococcus (Neotropical), Wapoacoccus Ben-Dov, 2007 Amonostherium – g/g Nipaecoccus (Australasian). Anisococcus – g/g Paraputo Annulococcus – g/g Boreococcus Deviating genera/species. Monotypic Anthelococcus – g/g Trionymus Albertinia, Antoninoides, Conicosoma, Para- Antonina – g/g Antonina serrolecanium as well as oligotypic Pseudan- Antoninella – g/g Mirococcopsis tonina save very small degenerate legs with Antoninoides – g/g Antonina partly fused segments. Monotypic genera Apodastococcus – g/g Pseudococcus Aemulantonina and Albertinia possess 1–2 Archanginella – g/g Metadenopus pairs of cerarii. Monotypic Miconicoccus Archeomyrmococcus – g/g Allomyrmococcus and Quadrigallicoccus produce galls. Asaphococcus – g/g Trionymus Asteliacoccus – g/g Trionymus Allomyrmococcus Atriplicicoccus – g/g Pseudococcus 19 g/g Takahashi, 1941 Atrococcus – g/g Pseudococcus = tribe Allomyrmococcini Williams, 1978 Australicoccus – g/g Nipaecoccus Benedictycoccina – g/g Rhizoecus Combination of diagnostic characters Bimillenia – g/g Mirococcopsis (apomorphic): both pairs of ostioles strong- Bolbococcus – g/g Allomyrmococcus ly developed, with sclerotized lips; unusu- Boninococcus – g/g Trionymus ally long, slender antennae and legs with Boreococcus – g/g Boreococcus long pointed claws and widely expanded Borneococcus – g/g Allomyrmococcus digitules; anal apparatus simplified, with- Brasiliputo – g/g Paraputo out pores and spinulae; very long dorsal and Brevennia – g/g Heterococcus ventral setae; cerarii and any conical setae Brevicoccus – g/g Rhizoecus absent; anal apparatus simplified, without Callitricoccus – g/g Trionymus pores and spinulae. Capitisetella – g/g Rhizoecus Chaetococcus – g/g Antonina Additional characters (plesiomorphic): Chaetotrionymus – g/g Trionymus claw without a denticle; trilocular pores nu- Chileputo – g/g Paraputo merous and evenly distributed. Chloeoon – g/g Antonina All genera of the group are endemic to Chlorococcus – g/g Trionymus Oriental region, mainly to Greater Sunda Chnaurococcus – g/g Trionymus Islands of Malay archipelago. Chorizococcus – g/g Pseudococcus Nominal genera (11): Allomyrmococcus Chryseococcus – g/g Pseudococcus Takahashi, 1941, Archeomyrmococcus Wil- Circaputo – g/g Paraputo liams, 2002, Bolbococcus Williams, 2002, Cirnecoccus – g/g Trionymus Borneococcus Williams, 2002, Dicranococcus Clavicoccus – g/g Pedronia Coccidella – g/g Rhizoecus Williams, 2002, Doryphorococcus Williams, Coccidohystrix – g/g Coccidohystrix 2002, Hippeococcus Reyne, 1954, Mala- Coccura – g/g Phenacoccus icoccus Takahashi, 1950, Paramyrmococcus Coleococcus – g/g Mirococcopsis Takahashi, 1941, Promyrmococcus Williams, Colombiacoccus – g/g Pseudococcus 2002, Thaimyrmococcus Williams, 2002. Conicosoma – g/g Antonina Conulicoccus – g/g Nipaecoccus Alphabetic index of generic names Coorongia – g/g Neotrionymus Cormiococcus – g/g Pseudococcus Acaciacoccus – g/g Mirococcopsis Crenicoccus – g/g Paraputo Acinicoccus – g/g Antonina Criniticoccus – g/g Paraputo Acrochordonus – g/g Pedronia Crisicoccus – g/g Trionymus Adelosoma – g/g Nipaecoccus Crocydococcus – g/g Trionymus Aemulantonina – g/g Antonina Cryptoripersia – g/g Mirococcopsis

Cucullococcus – g/g Mirococcus Lacombia – g/g Mirococcopsis Cyperia – g/g Paraputo Laingiococcus – g/g Heterococcus Cypericoccus – g/g Antonina Laminicoccus – g/g Paraputo Cypericoccus – g/g Metadenopus Lanceacoccus – g/g Nipaecoccus Cyphonococcus – g/g Nipaecoccus Lanceacoccus – g/g Paraputo Dawa – g/g Peliococcus Lankacoccus – g/g Paraputo Delococcus – g/g Paraputo Lantanacoccus – g/g Coccidohystrix Delottococcus – g/g Trionymus Lenania – g/g Mirococcopsis Dicranococcus – g/g Allomyrmococcus Leptococcus – g/g Trionymus Discococcus – g/g Mirococcus Leptorhizoecus – g/g Rhizoecus Distichlicoccus – g/g Pseudococcus Maconellicoccus – g/g Pseudococcus Diversicrus – g/g Mirococcopsis Macrocepicoccus – g/g Trionymus Doryphorococcus – g/g Allomyrmococcus Maculicoccus – g/g Trionymus Dysmicoccus – g/g Trionymus Madagasia – g/g Trionymus Eastia – g/g Puto Madangiacoccus – g/g Nipaecoccus Ehrhornia – g/g Mirococcus Madeurycoccus – g/g Mirococcopsis Epicoccus – g/g Nipaecoccus Malaicoccus – g/g Allomyrmococcus Erimococcus – g/g Peliococcus Malekoccus – g/g Phenacoccus Eriocorys – g/g Coccidohystrix Mammicoccus – g/g Phenacoccus Erioides – g/g Nipaecoccus Marendellea – g/g Trionymus Erium – g/g Trionymus Marottarhizoecus – g/g Rhizoecus Eucalyptococcus – g/g Paraputo Mascarenococcus – g/g Paraputo Eurycoccus – g/g Trionymus Mauricoccus – g/g Trionymus Exallomochlus – g/g Paraputo Melanococcus – g/g Nipaecoccus Exilipedronia – g/g Paraputo Metadenopus – g/g Metadenopus Extanticoccus – g/g Pedronia Miconicoccus – g/g Antonina Farinococcus – g/g Paraputo Mirococcopsis – g/g Mirococcopsis Ferrisia – g/g Trionymus Miscanthicoccus – g/g Neotrionymus Fijicoccus – g/g Nipaecoccus Misericoccus – g/g Metadenopus Fonscolombia – g/g Phenacoccus Mizococcus – g/g Mirococcopsis Formicococcus – g/g Paraputo Mollicoccus – g/g Mirococcus Gallulacoccus – g/g Pseudococcus Mombasinia – g/g Puto Geococcus – g/g Rhizoecus Moystonia – g/g Paraputo Glycycnyza – g/g Mirococcopsis Mutabilicoccus – g/g Paraputo Gomezmenoricoccus – g/g Mirococcopsis Nairobia – g/g Puto Grewiacoccus – g/g Nipaecoccus Natalensia – g/g Mirococcopsis Hadrococcus – g/g Mirococcopsis Neoclavicoccus – g/g Pedronia Hambletonrhizoecus – g/g Rhizoecus Neoripersia – g/g Neotrionymus Heliococcus – g/g Heliococcus Neosimmondsia – g/g Mirococcus Heterococcus – g/g Heterococcus Neotrionymus – g/g Neotrionymus Hippeococcus – g/g Allomyrmococcus Nesococcus – g/g Trionymus Hopefoldia – g/g Paraputo Nesopedronia – g/g Pedronia Hordeolicoccus – g/g Nipaecoccus Nesticoccus – g/g Antonina Humococcus – g/g Mirococcopsis Nipaecoccus – g/g Nipaecoccus Hypogeococcus – g/g Nipaecoccus Nudicauda – g/g Mirococcopsis Iberococcus – g/g Mirococcopsis Octococcus – g/g Phenacoccus Idiococcus – g/g Antonina Odacoccus – g/g Paraputo Inopicoccus – g/g Mirococcopsis Ohiacoccus – g/g Trionymus Ityococcus – g/g Trionymus Oracella – g/g Trionymus Kalaginella – g/g Metadenopus Orococcus – g/g Trionymus Kenmorea – g/g Nipaecoccus Orstomicoccus – g/g Trionymus Kermicus – g/g Antonina Oxyacanthus – g/g Phenacoccus Kissrhizoecus – g/g Rhizoecus Palaucoccus – g/g Mirococcopsis Lachnodiella – g/g Paraputo Palmicultor – g/g Paraputo

Paludicoccus – g/g Antonina Sphaerococcus – g/g Antonina Pandanicola – g/g Neotrionymus Spilococcus – g/g Pseudococcus Papuacoccus – g/g Pseudococcus Stemmatomerinx – g/g Phenacoccus Paracoccus – g/g Pseudococcus Stipacoccus – g/g Mirococcus Paradiscococcus – g/g Mirococcopsis Strandanna – g/g Trionymus Paradoxococcus – g/g Trionymus Stricklandina – g/g Paraputo Paraferrisia – g/g Trionymus Strombococcus – g/g Nipaecoccus Paramococcus – g/g Heterococcus Synacanthococcus – g/g Phenacoccus Paramonostherium – g/g Nipaecoccus Syrmococcus – g/g Metadenopus Paramyrmococcus – g/g Allomyrmococcus Tangicoccus – g/g Antonina Parapaludicoccus – g/g Antonina Tasmanicoccus – g/g Trionymus Paraporisaccus – g/g Antonina Telocorys – g/g Trionymus Paraputo – g/g Paraputo Thaimyrmococcus – g/g Allomyrmococcus Paraserrolecanium – g/g Antonina Tomentocera – g/g Paraputo Paratrionymus – g/g Metadenopus Trabutina – g/g Nipaecoccus Paulianodes – g/g Antonina Tridiscus – g/g Mirococcopsis Pedrococcus – g/g Pedronia Trimerococcus – g/g Puto Pedronia – g/g Pedronia Trionymus – g/g Trionymus Peliococcopsis – g/g Peliococcus Trochiscococcus – g/g Trionymus Peliococcus – g/g Peliococcus Turbinococcus – g/g Trionymus Pelionella – g/g Peliococcus Tylococcus – g/g Pedronia Peridiococcus – g/g Antonina Tympanococcus – g/g Trionymus Perystrix – g/g Phenacoccus Ventrispina – g/g Nipaecoccus Phenacoccus – g/g Phenacoccus Villosicoccus – g/g Trionymus Phyllococcus – g/g Trionymus Volvicoccus – g/g Mirococcopsis Pilococcus – g/g Mirococcopsis Wapoacoccus – g/g Antonina Planococcus – g/g Trionymus Williamsrhizoecus – g/g Rhizoecus Pleistocerarius – g/g Paraputo Yudnapinna – g/g Paraputo Poecilococcus – g/g Trionymus Porisaccus – g/g Antonina ACKNOWLEDGEMENTS Porococcus – g/g Nipaecoccus Promyrmococcus – g/g Allomyrmococcus The collection of Zoological Institute of Prorhizoecus – g/g Rhizoecus Russian Academy of Sciences was financially Prorsococcus – g/g Mirococcopsis supported by the Ministry of Education and Pseudantonina – g/g Antonina Science of the Russian Federation (project Pseudoferrisia – g/g Trionymus No. 16.518.11.7070). Pseudorhizoecus – g/g Rhizoecus Pseudoripersia – g/g Nipaecoccus REFERENCES Pseudotrionymus – g/g Trionymus Puto – g/g Puto Beardsley J.W. 1964. A new genus and species Pygmaeococcus – g/g Rhizoecus of Pseudococcidae from Haleakala, Maui. Quadrigallicoccus – g/g Antonina Proceedings of Hawaiian Entomological Soci- Rastrococcus – g/g Puto ety, 18(3): 365–368. Renicaula – g/g Trionymus Beardsley J.W. 1966. Homoptera: Coccoidea. Rhizoecus – g/g Rhizoecus Insects of Micronesia, 6: 377–562. Rhodania – g/g Metadenopus Beardsley J.W. 1971. New genera and species Ripersiella – g/g Rhizoecus of Hawaiian Pseudococcidae (Homoptera). Rosebankia – g/g Neotrionymus Proceedings of Hawaiian Entomological Soci- Sarococcus – g/g Paraputo ety, 21(1): 41–58. Scaptococcus – g/g Mirococcus Cox J.M. 1987. Pseudococcidae (Insecta: Hemip- Serrolecanium – g/g Antonina tera). Fauna of New Zealand, 11. Wellington. Seyneria – g/g Peliococcus 228 p. Sinococcus – g/g Mirococcus Danzig E.M. & Gavrilov-Zimin I.A. 2014. Pa- Spartinacoccus – g/g Trionymus laearctic mealybugs (Homoptera: Coccinea:

Pseudococcidae). Part 1. Subfamily Phe- and separation of higher taxa in Coccinea. nacoccinae. Fauna of Russia and neighbour- Zoosystematica Rossica, 22(1): 97–111. ing countries. New series, No. 148. Insecta: Gavrilov-Zimin I.A. & Matile-Ferrero D. 2014. Hemiptera: Arthroidignatha. St Petersburg: The genus Coccidohystrix Lindinger, 1943 Zoological Institute, Russian Academy of in the Palaeartic region with description of Sciences. 678 p. two new species from Maghreb (Homoptera: Danzig E.M. & Gavrilov-Zimin I.A. 2015. Pa- Coccinea: Pseudococcidae). Zoosystematica laearctic mealybugs (Homoptera: Coccinea: Rossica, 23(1): 96–105. Pseudococcidae). Part 2. Subfamily Pseudo- McKenzie H.L. 1967. Mealybugs of California coccinae. Fauna of Russia and neighbour- with taxonomy, biology and control of North ing countries. New series, No. 149. Insecta: American species (Homoptera: Coccoidea: Hemiptera: Arthroidignatha. St Petersburg: Pseudococcidae). Berkeley & Los Angeles: Zoological Institute, Russian Academy of Univ. California Press. 525 p. Sciences. 619 p. Millar I.M. 2002. Mealybug genera (Hemiptera: De Lotto G. 1957. The Pseudococcidae (Hom.: Pseudococcidae) of South Africa: identifica- Coccoidea) described by H.C. James from tion and review. African Entomology, 10(2): East Africa. Bulletin of the British Museum 185–233. (Natural History). Entomology, 5(5): 185– Mamet J.R. 1940. A new genus and five new 232. species of Coccidae (Hemipt. Homopt.) from De Lotto G. 1958. The Pseudoccoidae (Hom.: Mauritius. Proceedings of the Royal Entomo- Coccoidea) described by C.K. Brain from logical Society of London. Series B, Taxonomy, South Africa. Bulletin of the British Museum 9(4): 65–72. (Natural History). Entomology, 7(1): 79–120. Mamet J.R. 1951. Notes on the Coccoidea of De Lotto G. 1964. Observations on African Madagascar, II. Memoires de l’Institut Scien- mealy bugs (Hemiptera: Coccoidea). Bulletin tificue de Madagascar, 1951(5): 213–254. of the British Museum (Natural History). En- Mamet J.R. 1954. Notes on the Coccoidea of tomology, 14(8): 343–397. Madagascar, III. Memoires de l’Institut Scien- De Lotto G. 1967. The mealybugs of South Af- tificue de Madagascar, 1954(4): 1–86. rica (Homoptera: Pseudococcidae), I. Ento- Mamet J.R. 1962. Notes on the Coccoidea of mology Memoirs (South Africa), 12: 1–28. Madagascar, V (Homoptera). La Naturaliste De Lotto G. 1969. The mealybugs of South Af- Malgache Tananarive, 13: 153–202. rica (Homoptera: Pseudococcidae), II. Ento- Mamet J.R. 1967. New genera and species of Coc- mology Memoirs (South Africa), 20: 1–30. coidea from the Mascarene Islands (Homop- De Lotto G. 1977. On some African mealybugs tera). Mauritius Institute Bulletin, 6: 89–102. (Homoptera: Coccoidea: Pseudococcidae). Williams D.J. 1985. Australian mealybugs. Brit- Journal of the Entomological Society of South- ish Museum (Natural Hist.) special publica- ern Africa, 40: 13–36. tion No. 953. 431 p. Ferris G.F. 1948. Descriptions of new species, Williams D.J. 2004. Mealybugs of southern Asia. notes and figures. In: Zimmerman E.C. 1948. Kuala Lumpur. 896 p. Homoptera: Sternorrhyncha. Insects of Ha- Williams D.J. & Granara de Willink M.C. waii, 5: 141–276. 1992. Mealybugs of Central and South Ameri- Ferris G.F. 1950. Atlas of the scale insects of ca. London: CAB International. 635 p. North America. Ser. 5. V. 5. The Pseudococci- Williams D.J. & Watson G.W. 1988. The scale dae (Part 1). California. VII+278 p. Ferris G.F. 1953. Atlas of the Scale Insects of insects of the Tropical South Pacific Region. North America, V. 6. The Pseudococcidae Pt. 1. The Armoured Scales (Diaspididae). (Part II). California. 506 p. Wallingford: CAB International. 290 p. Gavrilov-Zimin I.A. & Danzig E.M. 2012. Ta- xonomic position of the genus Puto Signoret Received 28 October 2015 / Accepted 30 November 2015 (Homoptera: Coccinea: Pseudococcidae) Editorial responsibility: D.A. Gapon