COCCIDAE Fallén 1814

Hodgson, Chris, 2020, A review of neococcid scale insects (Hemiptera: Sternorrhyncha: Coccomorpha) based on the morphology of the adult males, Zootaxa 4765 (1), pp. 1-264 : 218-224

COCCIDAE Fallén 1814

Coccus Linnaeus 1758, 455 . Type species: Coccus hesperidum Linnaeus by subsequent designation.

Calymmata Costa 1828a, 6 . Type species: Coccus hesperidum Linnaeus. Type designation unknown. Junior synonym by community of type species.

Calypticus Costa 1829, 8 . Type species: Coccus hesperidum Linnaeus by subsequent designation Costa 1835. Junior synonym, Fernald 1903.

Type species: Coccus hesperidum Linnaeus.

Introduction. The family Coccidae is the third largest family in the Coccomorpha , with 169 genera and 1184 species ( García Morales et al. 2019). In almost all cladistic analyses, the Coccidae and Aclerdidae are sister families, and together are frequently sister to a clade containing the Asterolecaniidae , Kerriidae , Cerococcidae and Lecanodiaspididae (and occasionally the Kermesidae ) ( Gullan & Cook 2007; Hodgson & Hardy 2013; Vea & Grimaldi 2016). The classification within this large family was last revised by Hodgson (1994) who considered it was divided into 10 subfamilies, with the Coccinae including 4 tribes (Coccini, Paralecaniiini, Pulvinariini and Saissetiini). However, recent molecular studies (Kondo & Cook unpublished; Choi & Lee 2019) and the morphological study of the adult males by Hodgson and Hardy (unpublished) have cast doubt on this classification which suggest that the Coccidae might be non-monophyletic and should possibly include the Micrococcidae and Aclerdidae .

The first good description of an adult male soft scale was of Parthenolecanium (Eulecanium) pomeranicum (Kawecki) by Theron (1958), although he did not describe the setae. Shortly after that, Giliomee (1967a) described the males of a further 23 species belonging to 20 genera. Since then, the adult males of about another 90–100 species of soft scale insects have been described (see Appendix A ).

Family diagnosis based on adult male morphology ( Figs 87–94 ). Body: mostly small to moderate in size; body generally fairly setose, usually with both hs and fs; fleshy setae (fs) generally easy to differentiate from hair-like setae. Body rarely with small convex pores (cp). Head: median crest (mc) well developed and reticulated. Dorsal mid-cranial ridge short or absent; ventral mid-cranial ridge (vmcr) represented by a quite long medial ridge; with hs and/or fs ventral median ridge setae (vmcrs) posterior to each scape. Genae (g) usually unsclerotised, with or without polygonal reticulations; genal setae (gs) present or absent. Eyes: generally two pairs of simple eyes, subequal in size, rarely 3–5 when more lateral eyes sometimes smaller. Ocelli (o) present, usually placed close to postocular ridge. Ocular sclerite (ocs) sclerotised, covered in polygonal reticulations. Preocular ridge distinct both dorsally and ventrally. Postocular ridge (procr) generally well developed, occasionally not reaching ocelli. Postoccipital ridge generally absent. Interocular ridge rare. Antennae: 10 segmented (rarely 9 segmented), filiform. Thorax: prothorax: pronotal ridge (prnr) strong and often fused dorsally; without lateral pronotal setae; prosternum (stn 1) either without or with only a weak median ridge but with a strong transverse ridge; with or without hs prosternal setae. Small convex pores (cp) rare ( Poropeza ). Mesothorax: prescutal setae absent. Scutum (sct): median membranous area well developed, about twice as wide as long. Scutellar setae (scls) absent. Area within mesopostnotum unsclerotised. Basisternum (stn 2) with median ridge (mdr) either well developed or represented by weak sclerotisations medially; without basisternal setae (present on Milviscutulus ); lateropleurite (lpl) fairly large. Mesothoracic spiracle (sp 2) without disc-pores. Metathorax: with hs or fs metatergal seta (mts). Metathoracic spiracle (sp 3) without disc-pores. Metasternum (stn 3) lightly sclerotised. Small convex pores (cp) absent (except Poropeza ). Wings: hyaline; without alar sensoria and usually without alar setae. Hamulohalteres (h) present or absent, with generally 1 hamulus (rarely 2 or 3). Legs: generally quite setose, with both hs and fs; some setae spur-like. Tibia (ti): with 1 apical spur (tibs). Tarsi (ta) generally 1 segmented (2 segmented in Pounamococcus spp.) with tarsal spurs (tabs); tarsal campaniform pore (tcp) absent (present in Pounamococcus spp.); tarsal digitules (tdt) capitate. Claws (c) with or without a denticle; claw denticles capitate. Abdomen: caudal extension (ce) of segment VII generally well developed. Dorsal abdominal setae (ads) hs and/or fs; pleural setae (dps and vps) hs, fs or both; ventral abdominal setae (avs) often more abundant than dorsal abdominal setae. Segment VIII: caudal extension generally present but may be rounded with a cicatrix; ante-anal setae generally present; sternite (as) sclerotised, forming anterior margin to basal membranous area; without setae. Glandular pouches (gp) present or absent. Genital segments: penial sheath (ps) elongate and stout; segment IX completely fused to style, articulating anteriorly with sternite of segment VIII; narrowing to a blunt, rather parallel-sided narrow style. Anus (an) often obscure, without a strongly sclerotised margin. Basal membranous area distinct ventrally. Aedeagus (aed) quite short and broad, not nearly reaching apex of penial sheath.

The only classification that has been suggested to-date was entirely intuitive based on the morphology of the adult female ( Hodgson 1994) but taking into account the findings of Giliomee’s (1967a) study on males. As indicated above, neither molecular studies (T. Kondo and L.G. Cook, unpublished) nor that based on adult male morphology (Hodgson & Hardy, unpublished) have supported this classification.

Key to species of Coccidae based on adult male morphology (mainly based on published descriptions)

1. Tarsi 2 segmented. Tarsus with a tarsal campaniform pore. Fleshy setae absent on body. Glandular pouches absent. Hamulohalteres present................................................................. Pounamococcus Henderson & Hodgson

- Tarsi 1 segmented. Tarsus without a tarsal campaniform pore. Fleshy setae often present on body. Glandular pouches and hamulohateres present or absent........................................................................ 2

2. Two pairs of simple eyes present, a pair dorsally and another ventrally, and generally with a pair of ocelli............... 3

- Three or more pairs of simple eyes present and generally with a pair of ocelli.................................... 31

Species with only 2 pairs of simple eyes

3. Antennae 9 segmented................................................................. Myzolecaniinae … 4

- Antennae 10 segmented................................................................................ 7

Species with nine-segmented antennae

4. Interocular ridge present.............................................. Mesolecanium nigrofasciatum Pergande #

- Interocular ridge absent................................................................................ 5

5. Legs truncate. Tibia subequal in length to tarsus. Tibial spur about half length of tarsus (true of both macropterous and apterous forms)............................................................... Pseudophilippia quaintancii Cockerell

- Legs well developed. Tibia longer than tarsus. Tibial spur much less than half length of tarsus........................ 6

6. Antennal segments III–V together longer than length of penial sheath................. Neolecanium cornuparvum (Thro)

- Antennal segments III–V together subequal to or shorter than length of penial sheath …........ Toumeyella Cockerell spp.

Species with ten-segmented antennae

7. Postocular ridge long, extending well past each ocellus.....................................................… 8

- Postocular ridge short, either not nearly reaching each ocellus or only just reaching and then generally with an arm surrounding each ocellus …...................................................................................... 26

8. Fleshy setae entirely absent from body (but maybe present on antennae or legs) …................................. 9

- Fleshy setae present somewhere on body, often extremely abundant ….......................................... 16

9. Hamulohalteres absent ….............................................................................. 10

- Hamulohalteres present …............................................................................. 11

10. Most tergites and sternites sclerotised. Scutal and scutellar setae absent. Dorsal abdominal and ventral abdominal setae absent …........................................................... Cribropulvinaria tailungensis Hodgson & Martin

- Tergites and sternites membranous. Setae present on both scutum and scutellum. Dorsal abdominal and ventral abdominal setae present ….........................................................? Pulvinaria dodonaeae Maskell ǂ ( Fig. 94 )

11. Abdomen gradually narrowing from about segment IV to penial sheath, with almost no indication of caudal extensions on segments VII and VIII. Single alar seta present. Postmetaspiracular setae present. Mesepisternum reticulated ….......................................................................................... Cissococcus fulleri Cockerell

- Abdominal segments VII and VIII with distinct caudal extensions. Alar seta absent. Postmetaspiracular setae absent. Mesepisternum not reticulated …............................................................................. 12

12. Glandular pouches absent. Caudal extensions on abdominal segment VII rounded or absent; caudal extensions on abdominal segment VIII sclerotised (possibly a cicatrix), with five hs setae. Tegular setae absent..................................................................................................... Australian Akermes spp. ǂ ( Figs 88 , 89 )

- Glandular pouches present. Caudal extensions on abdominal segment VII distinct but short, those on abdominal segment VIII not sclerotised, with 2–4 hs setae. Tegular setae present...................................................... 13

13. Scutal setae present. Dorsal abdominal setae present on segments I–VI. Haltere with one hamulus.................... 14

- Scutal setae absent. Other characters not in this combination.................................................. 15

14. Compound eyes relatively small. Gena with few reticulations. Ocelli touching postocular ridge............................................................................................ Palaeolecanium bituberculatum (Targ.) .

- Compound eyes exceptionally large and bulging. Gena with reticulations throughout. Ocelli placed some distance from post- ocular ridge................................................. Fistulococcus pokfulamensis Hodgson & Martin

15. Tarsi very short and broad, less than half length of tibia and less than 3 times longer than wide. Dorsal abdominal setae absent on segments II–VI. Haltere often with 2 hamuli. Procoxal spurs absent.............. Physokermes Targioni Tozzetti spp.

- Tarsi quite long and narrow, about half as long as tibia and several times longer than wide. Dorsal abdominal setae present on segments II–VI. Haltere with 1 hamulus. Procoxal spurs stout and blunt......... Ctenochiton eucalypti Maskell ǂ ( Fig. 91 )

16. Caudal extensions entirely absent. Postmesospiracular setae present but restricted to a group posterior to each anterior spiracle. Membranous area of scutum with fleshy setae............................... Pseudopulvinaria sikkimensis Atkinson

- Caudal extensions well developed on at least abdominal segment VIII and generally on segment VII as well. Other characters not in this combination............................................................................... 17

17. Basisternal setae present...................................................... Milviscutulus mangiferae (Green)

- Basisternal setae absent............................................................................... 18

18. Interocular ridge present or postocular ridge fusing with preocular ridge. Fleshy setae present on some of abdominal segments III–VII............................................................................................ 19

- Interocular ridge absent. Postocular ridge never fusing with preocular ridge. Fleshy setae absent from dorsal surface of abdominal segments III–VII................................................................................. 22

19. Glandular pouches present............................................................................ 20

- Glandular pouches absent............................................................................. 21

20. Genal setae present. Prosternal setae present. Abdomen with both fs and hs dorsal abdominal setae............................................................................................ Luzulaspis caricis Ehrhorn ( Fig. 93 )

- Genal setae absent. Prosternal setae absent. Dorsal abdominal setae sparse and all hs............. Eriopeltis spp. Signoret

21. Postocular ridge extending to and fusing with preocular ridge just posterior to each scape... Psilococcus parvus Borchsenius

- Postocular ridge not fusing with preocular ridge but joined by a distinct interocular ridge just posterior to each scape.................................................................................... Luzulaspis luzulae (Dufour)

22. Postmesospiracular setae absent. Dorsal abdominal setae all hs, fleshy setae absent................................ 23

- Postmesospiracular setae present, extending across segment just anterior to basisternum. Dorsal abdominal setae generally including abundant fleshy setae......................................................................... 25

23. Glandular pouches absent. Genae with many fs setae. Prosternal setae absent. Legs well developed. Dorsospiracular setae pres- ent......................................................................... Hemilecanium Newstead spp.

- Glandular pouches present. Other characters not in this combination........................................... 24

24. Genal setae absent. Aedeagus short, about half length of penial sheath. Dorsal abdominal setae all hs and setae absent from segments II and III.................................................. Sphaerolecanium prunastri (Fonscolombe)

- Genal setae present. Aedeagus very short, a quarter or less length of penial sheath. Dorsal abdominal setae present on all segments, including segments II and III, with both fs and hs..................... Pulvinaria glacialis Gertsson & Hodgson

25. Scutal setae on membranous area of scutum abundant and mostly fleshy......................................................................... Ceroplastes Gray spp.; Coccus hesperidum Linnaeus ; Parasaissetia nigra Takahashi *

- Scutal setae on membranous area of scutum all hair-like or with hair-like setae more abundant than fleshy setae........... Parthenolecanium corni (Bouché) ; P. persicae (Fabricius) ( Fig. 87 ); Pulvinaria Šulc spp.; Pulvinariella mesembryanthemi (Vallot) and Prionococcus fontaniae Williams et al. *

26. All fleshy setae on antennae very short and blunt, each much shorter than width of antennal segments. Fleshy setae present on legs and antennae but restricted to a group of postmetaspiracular setae on body................................................................................................... Phalacrococcus howertoni Hodges & Hodgson

- Fleshy setae on antennae all much longer, each longer than width of antennal segments. Fleshy setae, if present on body, not restricted to postmetaspiracular setae.................................................................... 27

27. Small convex pores/simple pores frequent on dorsal surface of head, thorax and abdomen............................

.................................................................. Poropeza dacrydii Henderson & Hodgson - Small convex pores/simple pores, if present, absent from thorax and abdomen................................... 28

28. Simple pores present ventrally on head. Cicatrix present medially on dorsum of abdomen....... Inglisia patella Maskell #

- Simple pores absent from both surfaces of head. Cicatrix absent medially on dorsum of abdomen.................... 29

30. Apex of aedeagus spoon-shaped. Dorsospiracular setae absent. Tegular setae absent. Prosternal setae all hair-like setae.....

........................................................................... Crystallotesta fusca (Maskell) - Apex of aedeagus blunt but not spoon-shaped. Dorsospiracular setae present. Tegular setae present. Prosternal setae all fleshy setae.......................................................................... Lecanochiton Maskell spp.

With more than two pairs of simple eyes

31. Fleshy setae present over much of body, antennae and legs and often abundant................................... 32

- Fleshy setae on body either absent altogether or very few, generally restricted to a group of postmetaspiracular setae..... 36

32. Segmentation between trochanter and femur absent or indistinct, particularly on metathoracic leg. With some fleshy metatergal setae present. Penial sheath narrowing abruptly just prior to apex................... Plumichiton Henderson & Hodgson

- Segmentation between trochanter and femur clear and distinct. Metatergal setae rarely including fleshy setae. Penial sheath not narrowing abruptly but with apex generally more rounded................................................... 33

33. Postocular ridge reaching each ocellus and extending around it. Abdominal segment VIII unusually long.............. 34

- Postocular ridge not nearly reaching each ocellus. Abdominal segment VIII not unusually long........................ Aphenochiton Henderson & Hodgson spp.; Crystallotesta Henderson & Hodgson ( fagi -group spp.); Ctenochiton Maskell spp.; Epelidochiton Henderson & Hodgson spp.; Kalasiris perforata (Maskell) ; Umbonichiton Henderson & Hodgson spp.*

34. Gena without polygonal reticulations. Fleshy dorsal abdominal setae and fleshy dorsal pleural setae abundant on many segments. Tegular setae present........................................................................... 35

- Gena with polygonal reticulations. Fleshy dorsal abdominal setae and fleshy dorsal pleural setae infrequent or absent from most segments. Tegular setae absent............................................ Ctenochiton serratus Green ǂ ( Fig. 92 )

35. Dorsal abdominal setae including many fleshy setae. Ocular sclerite with polygonal reticulations almost fusing with those of median ridge dorsally....................................... Crystallotesta Henderson & Hodgson ( ornata -group)

- Dorsal abdominal setae very few and all hair-like setae. Ocular sclerite with polygonal reticulations poorly developed and not nearly reaching median ridge dorsally........................................... Inglisia theobromae (Newstead) ǂ -

36. Hamulohalteres absent................................................................................ 37

- Hamulohalteres present............................................................................... 40

37. Fleshy setae few, entirely absent from body although usually present on antennae and legs.......................... 38

- Fleshy setae present but restricted to a group of postmetaspiracular setae................... Phyllostroma myrtilli (Kalt.)

38. Gena lightly polygonally reticulated. Postocular ridge short, not reaching ocelli..... Aphenochiton inconspicuus (Maskell)

- Gena without reticulations. Postocular ridge longer, extending dorsally past ocelli................................. 39

39. Dorsal mid-cranial ridge absent. Tegular setae present................................ Parafairmairia Cockerell spp.

- Dorsal mid-cranial ridge long and well developed. Tegular setae absent.......... Cardiococcus major (Maskell) ǂ ( Fig. 90 )

40. With 3 pairs of simple eyes............................................................................ 41

- With 4 or 5 pairs of simple eyes........................................................................ 42

41. Simple eyes much enlarged, ventral pairs touching. Antennal setae subequal to or longer than length of respective antennal segments...................................................................... Philephedra Cockerell spp.

- Simple eyes not greatly enlarged and not touching. Antennal setae much shorter, no more than twice width of antennal segments.................................................................. Nemolecanium abietus Borchsenius

42. With 4 pairs of simple eyes............................................................................ 43

- With 5 pairs of simple eyes............................................................................ 45

43. A group of simple pores present posterior to pronotal sclerite on prothorax. Each haltere with 1 hamulus. Scutal setae present.

Ventral arms of preocular ridge almost meeting. Gena with polygonal reticulations............ Alecanochiton Hempel sp. - Simple pores absent. Each haltere with 2 hamuli. Other characters not in this combination.......................... 44

44. Dorsal and ventral simple eyes each more than 35 µm wide.......................... Eulecanium kunoense (Kuwana)

- Dorsal and ventral simple eyes each less than 25 µm wide............................. Rhodococcus Borchenius spp.

45. Setae on legs and antennae extremely long, length subequal to length of most antennal segments. Without alar setae........

.................................................................................. Ericerus pela (Chav.) - Setae on legs and antennae much shorter, never more than about 2x width of most antennal segments. With or without 1 or 2 alar setae.......................................................................................... 46

46. Scutal seta absent. Antennae short, less than half body length. Postmetaspiracular setae, if present, all hs. 1 or 2 alar setae pres-

ent........................................................................... Eulecanium Cockerell spp. - Scutal seta present. Antennae long, more than half body length. With both fs and hs postmetaspiracular. Alar setae absent.... .............................................................................. Filippia viburni (Signoret)

Note: *The males of these groups of taxa are morphologically very similar, differing in fairly small character-states, the taxonomic significance of which is uncertain. For species identification within these groups, see Giliomee (1967a) and Hodgson & Henderson (2004).

ǂ These species are not congeneric with the type species of the genus. In the case of the five species from Australia, the Australian Coccidae are in the process of being revised (Gullan & Hodgson, in prep.) and will be transferred to new genera once the revision is completed.

# Inglisia theobromae is not congeneric with I. patella , the type species of Inglisia ; likewise, Mesolecanium nigrofasciatum is not congeneric with M. nocturnum (Cockerell & Parrott) , the type species of Mesolecanium .

Figures

FIGURE 87. Parthenolecanium persicae (Fabricius). Macropterous male. (Coccidae). Where A=cranial apophysis, D=fleshy seta, E=hair-like seta, and R=dermal reticulations on ocular sclerite.

FIGURE 88. Akermes scrobiculatus (Maskell). Macropterous male. (Coccidae). Where R=dermal reticulations on ocular sclerite.

FIGURE 89. Akermes pinguis (Maskell). Macropterous male.(Coccidae).Where K=procoxa showing coxal bristles and R=dermal reticulations on ocular sclerite.

FIGURE 90. Cardiococcus major (Maskell). Macropterous male. (Coccidae). Where A=cranial apophysis and R=dermal reticulations on ocular sclerite.

FIGURE 91. Ctenochiton eucalypti Maskell. Macropterous male. (Coccidae). Where R=dermal reticulations on ocular sclerite.

FIGURE 92. Ctenochiton serratus Green. Macropterous male. (Coccidae). Where A=cranial apophysis, D=fleshy seta, E=hairlike seta and R=dermal reticulations on ocular sclerite.

FIGURE 93. Luzulaspis caricis (Ehrhorn). Macropterous male. (Coccidae). Where A=cranial apophysis, and R=dermal reticulations on ocular sclerite.

FIGURE 94.?Pulvinaria dodonaeae Maskell. Macropterous male. (Coccidae). Where A=cranial apophysis and R=dermal reticulations on ocular sclerite.

Appendix A. A list of descriptions of extant adult male Coccomorpha, mainly since Theron’s pioneering work of 1958. It is unlikely to be complete, and so please contact the author if you know of others.