I have followed the classification that was suggested by Hodgson (1994) when he rediscribed the adult females of the type species of all the known soft scale genera. This classification was based on the structure of the adult females and on male morphology as studied since about 1960. He divided the Coccidae into 10 subfamilies, namely CARDIOCOCCINAE Hodgson, CEROPLASTINAE Atkinson, CISSOCOCCINAE Brain, COCCINAE Fallen, CYPHOCOCCINAE Hodgson, EULECANIINAE Koteja, ERIOPELTINA Šulc, FILIPPIINAE Bodenheimer, MYZOLECANIINAE Hodgson, PSEUDOPULVINARIINAE Tang, with the Coccinae divided into four tribes, viz., Coccini Fallén, Paralecaniini Williams, Pulvinariini Targoni Tozzetti and Saissetini Hodgson. He considered that the status of these groupings needed further study.
The family Coccidae is the third largest family within the Coccoidea, with approximately 1100 species in about 160 genera (Ben-Dov, 1993; Hodgson, 1994). A typical adult female soft scale is characterised by the presence of:
(i) a pair of more or less triangular anal plates (except in Physokermes),
(ii) a deep anal cleft,
(iii) sunken ventral microducts,
(iv) an eversible anal tube to assist in honeydew elimination,
(v) an anal ring composed of two sclerotised crescents, each with setae and pores
(vi) eyespots which, even when lying in the line of the marginal spines, appear to be slightly displaced onto the dorsum,
(vii) one-segmented labium,
(viii) marginal setae,
(ix) stigmatic spines (although sometimes absent),
(x) pregential multilocular disc-pores on the venter of the abdomen,
(xi) a band of spiracular disc-pores between the marginal and each spiracle,
(xii) ventral tubular ducts, frequently present in a submarginal band,
(xiii) legs and antennae usually well developed, and
(xiv) only three setae on scape.
(xv) absence of a campaniform pore on the tarsus
(xvi) absence of microtubular ducts (except in the Cyphococcinae),
(xvii) absence of true figure-of-eight pores (except in Bodenheirmera and Mallococcus where they are present).
(i) adult females of most genera produce some sort of test, which may be woolly (as in the Filippinae), glassy (as in the Cardiococcinae), thin wax (as in the Coccinae) or thick wax (as in the Ceroplastinae),
(ii) the dorsum of the mature adult females of many species becomes heavily sclerotised at maturity, and
(iii) the dorsum usually has dorsal setae, dorsal microductules and, frequently, preopercular pores, dorsal tubercles and tubular ducts.
The other classification was suggested by Hodgson (1994) when he rediscribed the adult females of the type species of all the known soft scale genera. This classification was based on the structure of the adult females and on male morphology as studied since about 1960. He divided the Coccidae into 10 subfamilies (Cardiococcinae, Ceroplastinae, Cissococcinae, Coccinae, Cyphococcinae, Eulecaniinae, Eriopeltinae, Filippiinae, Myzolecaniinae, Pseudopulvinariinae) with the Coccinae divided into four tribes (Coccini, Paralecaniinae, Pulvinariini, Saissetiini).
This subfamily, which might be loosely called the glassy scales, includes 16 genera. The adult females are characterised by:
(i) the presence of a glassy test,
(ii) the absence of dorsal setae, dorsal tubular ducts and dorsal tubercles,
(iii) presence of spinose marginal setae,
(iv) a distinctive distribution of dorsal pores, which occur in a distinctive pattern, often in a mid-dorsal line from the anal plates to the anterior margin of the head or forming a large reticulate pattern,
(v) presence of pregenital disc-pores with typically five loculi,
(vi) presence of a submarginal band of ventral tubular ducts
(vii) the inner margins of the anal plates often diverging posteriorly, with spinose setae along the inner margin in many genera, and
(viii) absence of pairs of long setae medially on the pregenital segments, these replaced by bands of rather spinose setae.
This subfamily includes all the genera related to Ceroplastes Gray. This group of genera is very distinctive and the adult females can be recognised by the presence of:
(i) a thick waxy test covering the dorsum,
(ii) the distinctive dorsal Ceroplastes-type pores,
(iii) a sclerotised caudal process, which lifts the anal plates above the thick wax cover,
(iv) the prsesnce of dorsal lobes or clear areas free from pores,
(v) the form of the ventral microducts, which appear to have a cruciform opening
(vi) the characteristics of the stigmatic areas.
(i) the reduction of the dorsum to a small area around the anal plates and their associated sclerotisations, the median areas of the venter becoming expanded, so that the legs, spiracles and mouthparts lie dorsally,
(ii) possible complete absence of antennae,
(iii) reduced legs,
(iv) large spiracles, placed on the apparent dorsum with their atria facing medially, each with sparse bands of five-locular disc-pores also extending medially,
(v) mouthparts also on the apparent dorsal surface, with the labium pointing anteriorly,
(vi) presence of anal plates typical of the Coccidae, with numerous setae on their dorsal surface (as in the Myzolecaniinae),
(vii) anal plates surrounded by a very large area of sclerotisation, which appears to be structurally quite different to that of other Coccidae,
(viii) true venter covered in numerous long setae and 10-locular disc-pores, and
(ix) absence of marginal setae and stigmatic spines.
Hodgson (1994) included 55 genera in four tribes in this subfamily. The tribes were as follows:
Members of this tribe are typically characterised by:
(i) lack of dorsal tubular ducts (except very sparsely submarginally in Coccus),
(ii) absence of ventral tubular ducts or their restriction to medially in the thorax,
(iii) lack of pocket-like sclerotisations,
(iv) presence of eyespots, usually close to the margin,
(v) stigmatic areas unsclerotised,
(vi) presence of stigmatic spines differentiated from the marginal setae,
(vii) presence of pregenital disc-pores concentrated on the pregenital segment, never present medially on the thorax or head.
Typical members of this tribe have
(i) a distinct cleft which is sclerotised on the dorsum at its base,
(ii) no dorsal tubular ducts,
(iii) ventral tubular ducts, when present, restricted to a group on either side of the genital opening,
(iv) pregenital disc-pores restricted to the abdominal segments immediately anterior to the genital opening,
(v) eyespots displaced onto the dorsum, typically nearly dorsal to the base of each antenna.
Members of this tribe have
(i) a woolly ovisac secreted by the reproducing female from beneath the posterior end of the abdomen, often lifting the insect so that its body is held almost vertically above the head by the ovisac,
(ii) ventral tubular ducts of generally three or four types (rarely two), including (a) a small duct with a fine inner ductile, generally occurring in a submarginal band, and (b) a larger duct with the inner and outer ductules of subequal width, these typically present medially in the head and thorax but occasionally elsewhere,
(iii) no woolly test covering the dorsum, (or, if a mealy covering is present, this is very sparse),
(iv) no dorsal tubular ducts, or if present, of one type only and typically similar to the smallest ventral ducts,
(v) spinose dorsal setae,
(vi) each leg with a tibio-tarsal articulation,
(vii) no pocket-like sclerotisations,
(viii) eyespots present near the margin, and
(ix) shallow, unsclerotised stigmatic clefts
Members of this tribe differ from other members of the Coccinae in:
(i) presence of a broad submarginal band of ventral tubular ducts of one or two types,
(ii) absence (typically) of dorsal tubular ducts,
(iii) typically with dorsal tubercles and, often, also pocket-like sclerotisations, though both may be absent,
(iv) presence of pregenital disc-pores, each usually with 10 loculi, extending medially onto thorax,
(v) presence of eyespots near the margin, and
(vi) presence of unsclerotised, shallow stigmatic clefts.
(i) presence of microtubular ducts on the dorsum, otherwise unknown in the Coccidae,
(ii) dorsum divided into two areas, the median area with few pores and no dorsal setae, the lateral areas with abundant pores and setae,
(iii) these two areas separated by a sinuous line of strongly spinose setae,
(iv) presence of pregenital disc-pores, each with six or seven loculi, frequent medially in all the abdominal and thoracic segments,
(v) absence of preopercular pores,
(vi) spiracular disc-pores present in broad bands,
(vii) presence of spinose marginal setae,
(viii) presence of tibio-tarsal pseudo-articulations,
(ix) claw digitules both fine, (x) absence of eyespots, and
(xi) presence of a glassy test covering the median area of the dorsum.
(i) pregenital disc-pores, each with 10 loculi, present medially on all the abdominal and thoracic segments and usually also on the head,
(ii) presence of spinose or setose marginal setae, which are never fimbriate,
(iii) absence of dorsal tubercles and pocket-like sclerotisations,
(iv) typically with a complete ring of ventral tubular ducts present,
(v) legs without a tibio-tarsal articulatory sclerosis, and
(vi) even though the legs are well developed, the claw digitules are either both fine or dissimilar, never both broad.
(i) body elongate,
(ii) production of a felted ovisac over the whole or part of the dorsum, which is secreted by
(iii) large tubular ducts on the dorsum, which are similar to the tubular ducts found submarginally on the venter,
(iv) a membranous dorsum, without areas of dense sclerotisation,
(v) each anal plate frequently with one or two setose or spinose setae along the inner margin,
(vi) lack of stigmatic clefts,
(vii) pregenital disc-pores each with 7-10 loculi,
(viii) presence typically of two types of ventral tubular ducts, the larger submarginally,
(ix) legs and antennae well developed,
(x) presence of either 0 or two stigmatic spines in each stigmatic area, and
(xi) lack of dorsal tubercles and pocket-like sclerotisations. In addition, members of this subfamily are usually restricted to monocotyledonous plants.
(i) The Filippiinae mainly occur on dicotyledonous plants
(ii) The adult females are roundly oval in shape, sometimes have dorsal tubercles and pocket-like sclerotisations and
(iii) They have 0, one, two or three stigmatic spines in each stigmatic cleft.
(iv) Otherwise they share most of the characters given above for the Eriopeltinae.
The main characters of the adult females are:
(i) the lack of dorsal tubular ducts,
(ii) absence of eyespots,
(iii) presence of anal plates with typically numerous setae on the dorsal surface,
(iv) particularly large and often somewhat modified spiracles (Hodgson, 1995), with broad bands of spiracular disc-pores between the margin and the spiracle,
(v) ventral tubular ducts of one type, frequently restricted to a group on either side of the genital opening,
(vi) without median pairs of long pregenital setae but with segmental bands of short spinose setae,
(vii) legs reduced, with both claw digitules fine,
(viii) reduced antennae, and
(ix) a short anal tube.
(i) the production of a dense, woolly test which covers the entire insect
(ii) abundant sclerotised, five-locular disc-pores or cribriform plates which cover the entire dorsum and also form a narrow submarginal band ventrally,
(iii) anal plates which appear to be joined along both the dorsal and ventral margins, each plate rather triangular in shape, with long spinose setae, appearing rather like a crown,
(iv) tube either very short or absent,
(v) extremely large spiracles,
(vi) lacking tubular ducts, but
(vii) with one or two types of tubular duct ventrally,
(viii) presence of strongly spinose marginal setae, but
(ix) with the stigmatic spines undifferentiated and
(x) legs and antennae more or less well developed
Ben-Dov, Y. 1993. A Systematic Catalogue of the Soft Scale Insects of the World (Homoptera: Coccoidea: Coccidae) with data on geographical distribution, host plants, biology and economic importance. Flora and Fauna Hnadbook No. 9. Sandhill Crane Press, Inc., Gainesville, Florida, xxviii + 536 pp.
Hodgson, C.J. 1994. The Scale Insect Family Coccidae: An Identification Manual to Genera. CAB International, Wallingford. Vi + 639 pp.
Hodgson, C.J. 1995. A brief review of the structure of the spiracle in the family Coccidae. Israel Journal of Entomology, 29: 47-55.
Hodgson, C.J. 1997. Classification of Coccidae and Coccid families. In: Soft Scale Insects: Their Biology, Natural Enemies and Control. (Eds. Ben-Dov, Y and Hodgson, C.) pp.157-201.