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- ditions 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- © 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 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