NEOCOCCOID

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 : 19-22

KEY TO EXTANT NEOCOCCOID HIGHER TAXA (MAINLY FAMILIES AND SUBFAMILIES) BASED ON ADULT MALE MORPHOLOGY

Macropterous and most brachypterous males.

(Note: this key includes all neococcoid families except Halimococcidae , Micrococcidae , Phoenicococcidae and Xenococcidae, the known males of which are all apterous – see key for apterous males below).

1. Compound eyes present. Abdominal spiracles generally present on at least some segments, each generally quite obvious, with a clearly sclerotised atrium or peritreme. Wings without microtrichia. Glandular pouches on abdominal segment VIII absent. Post-tergites, when present, generally lying more or less longitudinally along prothorax. Triangular plate absent........................................................................................... ARCHAEOCOCCOIDS

- Compound eyes absent, replaced by various arrangements of small, simple unicorneal eyes. Abdominal spiracles present (taxa close to Phenacoleachia ) or absent; when present, atrium or peritreme extremely difficult to detect, presence generally indicated by branching tracheae in some or all abdominal segments. Wings with microtrichia. Frequently with a pair of glandular pouches laterally on abdominal segment VIII. Post-tergites, when present, lying transversely, parallel to anterior margin of scutum/prealare. Triangular plate present................................................................... 2

2. Simple, unicorneal eyes in a band 1, 2 or more eyes wide around head, each ocular sclerite generally with 5 or more simple eyes. Scutum without a quadrate membranous area lying dorsally between prescutum and scutellum............................................................................... Phenacoleachia , Puto , Steingelia , and Pityococcus

- Simple, unicorneal eyes present either in a band only 1 eye wide, each ocular sclerite generally with 4 or fewer simple eyes or, most frequently, with only a dorsal pair and a ventral pair. If 4 or 5 pairs of simple eyes present ( Kermesidae and some Coccidae ), then scutum with a quadrate membranous area between prescutum and scutellum dorsally … NEOCOCCOIDS.... 3

3. Scutum without a distinct, quadrate membranous area between prescutum and scutellum, although some areas of scutum may be membranous. Ocular sclerite and genae without polygonal reticulations (although with striations or concentric ridges often present around simple eyes)............................................................................. 4

- Scutum with a distinct, quadrate membranous area between prescutum and scutellum, with lateral margins generally demarcated by narrow sclerotised ridges. Ocular sclerite and genae with polygonal reticulations (though sometimes modified).. 16

Scutum without a distinct, quadrate membranous area between prescutum and scutellum

4. Bifurcated setae never present on tarsi. Penial sheath of various shapes but rarely broad basally and then narrowing abruptly to a long, narrow shaft, 3 or more times longer than broad basal section............................................ 5

- Bifurcated setae present on tarsi (these may be hard to see on some specimens). Penial sheath very long, broad basally and then narrowing abruptly to a long, narrow shaft, 3 or more times longer than broad basal section......................... 15

5. Claw digitules setose. Claw long, narrowing gradually to a sharp apex. Antennae with capitate setae restricted to apical segment. Postoccipital ridge absent or poorly developed, and then slender, V- or U-shaped. Lateropleurite narrow. Wing sensoria and alar setae often present. Trochanter with a Y-shaped ridge distally. Glandular pouches, when present, with more than 2 setae............................................................................................... 6

- Claw digitules capitate. Claw broad at base, abruptly tapering towards apex (except Dactylopius ). Antennae often with capitate setae on all or most flagellar segments. Postoccipital ridge, when present, strong and laterally forked. Lateropleurite broad. Wing sensoria and alar setae generally absent. Trochanter without a Y-shaped ridge distally. Glandular pouches, when present, with only 2 setae..................................................................................... 8

6. Head with a pair of ocelli in addition to two pairs of simple eyes. Postocular ridge generally distinct. Postoccipital ridge generally distinct. Dorsal and ventral mid-cranial ridges usually well developed. Neck well developed between head and prothorax. Pronotal ridge and pronotal sclerite present. Propleural ridge well developed as a proepisternum + cervical sclerite which extends anteriorly to reach postocular ridge. Metaprecoxal ridge well developed. Ostioles between abdominal segments VI and VII often present. Glandular pouches usually present on abdominal segment VIII and occasionally also on segment VII. Generally macropterous, although occasionally brachypterous or apterous … PSEUDOCOCCIDAE Westwood. ......... 7

- Head without ocelli, although two pairs of simple eyes generally present. Postocular ridge absent although a faint line occasionally present marking posterior margin of ocular sclerite. Dorsal mid-cranial ridge absent. Ventral mid-cranial ridge poorly developed or absent. Neck entirely absent. Pronotal ridge and pronotal sclerite absent. Propleural ridge well developed but short, only extending anteriorly about half-way to head. Metaprecoxal ridge absent. Ostioles absent. Glandular pouches absent, although 1 or more long setae may be present in this position on abdominal segment VIII. Possibly mostly apterous or brachypterous, occasionally macropterous........................................... RHIZOECIDAE Williams ( Figs 2–5 )

7. With two pairs of glandular pouches, one pair on abdominal segment VII and other on VIII (only 1 pair on Rastrococcus spp). Preocular ridge ventrally long and distinct. Loculate pores usually present dorsally and along margins of abdomen in addition to those associated with glandular pouches. Lateropleurite broad. Penial sheath with a distinct constriction about half-way along length. Penial sheath without a distinct ventral slit extending to basal rod............ PHENACOCCINAE Šulc ( Figs 8 , 9 )

- Either without glandular pouches or with only a single pair on segment VIII. Preocular ridge absent ventrally. Loculate pores mainly restricted to abdominal margins and glandular pouches. Lateropleurite narrow. Penial sheath tending to narrow gradually, without a distinct constriction about half-way along length. Penial sheath with a distinct ventral slit extending to basal rod............................................................ PSEUDOCOCCINAE Cockerell ( Figs 10–13 )

8. Sensilla on each side of trochanter roundish and more or less forming a triangle. Antennae with 10 segments............ 9

- Sensilla on each side of trochanter rather oval to elongate and more or less in a straight line. Antennae with 9 or fewer segments............................................................................................. 12

-

9. Glandular pouch setae very short, less than 1/4 length of penial sheath, each with a blunt apex. Postoccipital ridge absent. Interocular ridge present. Claw long and narrow. Interocular ridge present.......... DACTYLOPIIDAE Costa ( Figs 24 , 25 )

- Glandular pouch setae, when present, much longer and setose. Postoccipital ridge well developed, with lateral anterior and posterior extensions. Claw broad at base, abruptly tapering towards apex. Interocular ridge absent..................... 10

10. Hamulohalteres usually present (except Ovaticoccus ). Postmesospiracular setae present or absent. Capitate setae present on at least some antennal segments.......................................................................... 11

- Hamulohalteres absent. Postmesospiracular setae absent. Capitate setae absent on all antennal segments....................................................................................... Apiomorpha Rübsaamen ( Figs 19–23 )

11. Alary setae absent. Postmesospiracular setae present. Capitate setae restricted to apical antennal segment................................................................................... Eriococcus buxi (Fonscolombe) ( Fig. 42 )

- Alary setae present. Postmesospiracular setae absent. Capitate setae present on other antennal segments in addition to apical segment.. ACANTHOCOCCIDAE GROUP ( Acanthococcus Signoret ; Eriochiton Maskell ; Gossyparia Signoret ; Ovaticoccus Kloet ) ( Figs 15–18 )

12. Antennae each with 9 segments. Penial sheath extremely long and needle-like, length greater than half total body length....

................................................................ Cylindrococcus spiniferus Maskell ( Fig. 43 ) - Antennae with 8 or fewer segments. Penial sheath not needle-like and never as long as half total body length........... 13

13. Antennae 8 or more segmented................. ERIOCOCCIDAE ‘GONDWANA GROUP’ ( Callococcus Ferris ( Fig. 26 ); Calycicoccus merwei Brain ( Fig. 27 ); Capulinia Signoret ( Fig. 28 ); Carpochloroides Cockerell ( Fig. 29 ); Choneochiton Hodgson ( Fig. 30 ); Cystococcus Fuller ( Fig. 31 ); Dromedaricoccus Hodgson & Miller ( Fig. 32 ); Eriogallococcus Hodgson & Magalhãaes ( Fig. 33 ); Lachnodius Maskell ( Fig. 34 ); Lobimargo Hardy & Gullan ( Fig. 35 ); Pseudotectococcus rolliniae Hodgson & Gonçalves ( Fig. 37 ); Stibococcus Miller & Gonzalez ( Fig. 38 ); Tanyscelis Hardy & Gullan ( Fig. 39 ); Tectococcus Hempel ( Fig. 40 ).

- Antennae with 7 or fewer segments.......................................................................... 14

14. Antennae seven segmented. Tarsal digitules normal, both arising more or less on dorsal surface of tarsus. Penial sheath divided into anterior segment IX and posterior style, but without a distinct “segmental” membrane separating them.................. BEESONIIDAE Green ( Beesonia Green ; Danumococcus Takagi ; Echinogalla Takagi ; Gallacoccus Takagi ; Mangalorea Takagi ) ( Figs 44–46 )

- Antennae 6 or 7 segmented. Tarsal digitules unusual, with one arising as normal from dorsal surface, other arising from close to base of claw laterally. Tarsal digitules often dissimilar in shape. Penial sheath clearly divided into anterior segment IX and posterior style, separated by a “segmental” membrane.................... STICTOCOCCIDAE Cockerell ( Figs 47–52 )

Scutum without a distinct, quadrate membranous area between prescutum and scutellum; tarsi with bifurcated

setae

15. Antennae 10-segmented. Postoccipital ridge large and well defined. Tibio-tarsal articulation distinct. Abdominal segment IX not clearly separate from penial sheath. Claw long and narrow, without a denticle. Lateropleurite narrow. Trochanters particularly long, usually about 1/3rd length of trochanter + femur. Apical antennal segment pointed, apex with a single capitate seta. Ocular sclerite unsclerotised................................................ DIASPIDIDAE Costa ( Figs 59–61 )

- Antennae with 9 or fewer segments. Postoccipital ridge absent. Tibia and tarsus fused, without any articulation. Abdominal segment IX clearly seperate from penial sheath, forming a distinct segment. Claw broad basally, with a distinct denticle. Lateropleurite broad. Trochanters not exceptionally long, each about 1/4th length of trochanter + femur. Apical antennal segment with a rounded apex and with 4 capitate setae. Ocular sclerite showing some sclerotisation.......................................................................................... CONCHASPIDIDAE Cockerell ( Figs 53–56 )

Scutum with a distinct, quadrate membranous area between prescutum and scutellum.

16. Setae on margins of penial sheath mainly quite long, each more than 10 µm long................................. 17

- Setae on margins of penial sheath all minute, mainly less than 5 µm long........................................ 20

17. Membranous area of mesothorax posterior to scutellum with polygonal reticulations throughout. Membranous area of scutum (between prescutum and scutellum) also often polygonally reticulated. Ventral surface of claw S-shaped, with broad basal part narrowing abruptly to a long, narrow apical part without a denticle. Tibia without tibial spurs. Segment VIII of abdomen present as an almost complete sclerotised ring........................ ASTEROLECANIIDAE Targioni Tozzetti ( Figs 70–74 )

- Membranous area of mesothorax posterior to scutellum without polygonal reticulations. Membranous area of scutum (between prescutum and scutellum) without polygonal reticulations. Ventral surface of claw either not S-shaped as above or, if rather S-shaped, then with a denticle. Tibia with 1 or 2 tibial spurs. Segment VIII of abdomen, if sclerotised, not forming a complete sclerotised ring...................................................................................... 18

18. With 5 pairs of simple eyes, more or less arranged in a ring around head. Hamulohalteres present. Preocular ridge fusing medially or nearly so, at least extending over top of ventral simple eye. Penial sheath narrowing abruptly between segment IX and style. Cranial apophysis bifurcated......................................... KERMESIIDAE Boitard ( Figs 62–64 )

- With only 2 pairs of simple eyes, one pair dorsally and other ventral. Hamulohalteres absent on known species. Preocular ridge never fusing medially and only rarely extending over top of ventral simple eye. Penial sheath not narrowing abruptly between segment IX and style. Cranial apophysis not bifurcated...................................................... 19

19. Penial sheath broad basally, narrowing abruptly about 1/4 along length to a long, thin, narrow style. Fleshy setae present on venter of abdomen. Ventral abdominal setae many times more abundant than dorsal abdominal setae. Tibia with two tibial spurs. Genae with polygonal reticulations. Glandular pouch setae, when present, subequal in length. Each glandular pouch deep................................................................... KERRIIDAE Targioni Tozzetti ( Figs 65 , 66 )

- Penial sheath more or less triangular, not narrowing abruptly about 1/4 along length to a long thin narrow style. Fleshy setae absent on venter of abdomen. Ventral abdominal setae as abundant as dorsal abdominal setae. Tibia with one tibial spur. Genae without polygonal reticulations. Glandular pouch setae, when present, with one seta about half length of other. Each glandular pouch shallow..................................................... CEROCOCCIDAE Comstock ( Figs 67–69 )

20. Membranous area of mesothorax posterior to scutellum usually with a large area of sclerotisation. Penial sheath short, length generally less than 3x greatest breadth. Sternite VIII sclerotised, with a sclerotised ridge along lateral margins. Thoracic spiracles each with associated locular pores. Anus with a strongly sclerotised margin. Caudal extensions to abdominal segment VIII absent................................................................ ACLERDIDAE Signoret ( Figs 78–86 )

- Membranous area of mesothorax posterior to scutellum entirely membranous. Penial sheath long, length generally more than 4x greatest breadth. Sternite VIII generally without a sclerotised ridge along lateral margins. Thoracic spiracles without associated locular pores. Anus with membranous margins. Caudal extensions to abdominal segment VIII present, if only rounded. .................................................................................................. 21

21. Trochanter and femur fused (forming a trochantofemur). Tarsi with bifurcate setae. Preocular ridge, if present, extremely short. Ocelli present. With only 2 pairs of simple eyes. Basal membranous area at anterior end of penial sheath small or absent ventrally. Postoccipital ridge present. Glandular pouches absent. Aedeagus long, reaching or nearly reaching apex of penial sheath................................................................ LECANODIASPIDIDAE (Figs 75,76)

- Trochanter and femur not fused. Tarsi without bifurcate setae. Preocular ridge always well-developed both dorsally and ventrally. Ocelli absent. Occasionally with up to 5 pairs of simple eyes. Basal membranous area present at anterior end of penial sheath. Postoccipital ridge generally absent. Glandular pouches present or absent. Aedeagus shorter, not nearly reaching apex of penial sheath.......................................................... COCCIDAE Linnaeus ( Figs 87–94 )

Apterous and a few brachypterous adult males.

Note: apterous adult males are known in the Eriococcidae , Halimococcidae , Micrococcidae , Phoenicococcidae , Pseudococcidae , Rhizoecidae and Xenococcidae. A few brachypterous males are known with very reduced or no thoracic sclerotisation, e.g., the pseudococcid Mirococcus (Polystomophora) ostiaplurimus (Kiritchenko, 1940) .

1. Compound eyes present............................................................... ARCHAEOCOCCIDS

- Compound eyes absent................................................................................ 2

2. Simple eyes and ocelli absent........................................................................... 3

- Simple eyes and/or ocelli present........................................................................ 4

- Legs very much reduced, anterior legs 1 segmented, meso- and metathoracic legs 3 or 4 segmented. Antennae reduced to a single segment with a bunch of about 20 setae. Each caudal extension on abdominal segment VIII unsclerotised and rounded. Dermal pores entirely absent. Penial sheath short and stout, length subequal to basal width.................................................................................... ACLERDIDAE ) ( Nipponaclerda McConnell , Fig. 85 )

4. Claw digitules apically clubbed. Claws not long and narrow, rather broad basally and quite short. Simple eyes present or absent but ocelli present. Tarsi only one segmented. Loculate pores absent. Antennal segment III very narrow (stalk-like) basally.. 5

- Claw digitules short and setose. Claws generally long and narrow, tapering to a sharp point. With or without glandular pouches, when pouches absent, generally with a few longer setae in this position. Simple eyes generally present. Tarsi 1 or 2 segmented. Loculate pores generally present. Antennal segment III not especially narrow and stalk-like.......................... 7

5. Abdomen at least as long as thorax and tapering evenly to penial sheath. Penial sheath triangular and about as long as previous segment. Tibial spurs present. Abdominal tergites and sternites absent................................................................................................ CRYPTOCOCCIDAE ( Pseudochermes Cockerell ) ( Fig. 41 )

- Abdomen short, often only as long as metathorax. Penial sheath of variable shape but much longer than previous segment. Tibial spurs absent. Abdominal tergites and sternites generally present, at least on segment VIII....................... 6

6. Antennae rather short, with distal 4 segments forming a club. Apical antennal segment with a knob-like apex. Fleshy setae on antennae short. Legs rather reduced. Tarsi with sensory areas. Tarsus subequal in length to tibia. Fleshy setae present only on apical 3 antennal segments. Fleshy setae absent from legs.................. PHOENICOCOCCIDAE Cockerell ( Fig. 57 )

- Antennae short or long, but either with only distal 3 antennal segments forming a club or without a club. Apical antennal segment without a knob-like apex. Fleshy setae on antennae quite long. Tarsi without sensory areas. Fleshy setae present on all antennal segments. Fleshy setae present on legs, at least femora............... HALIMOCOCCIDAE Cockerell ( Fig. 58 )

7. Head with two pairs of simple eyes plus a pair of ocelli. Postocular ridge distinct. Postoccipital ridge present or absent. Ventral mid-cranial ridges usually well developed. Propleural ridge generally well developed as a proepisternum + cervical sclerite, extending anteriorly to reach head. Ostioles sometimes present between abdominal segments VI and VII. Glandular pouches usually present on abdominal segment VIII, each with a group of loculate pores....... PSEUDOCOCCIDAE ( Figs 13 , 14 )

- Head without ocelli, although two pairs of simple eyes present on most Rhizoecinae . Postocular ridge absent although a faint line occasionally present marking posterior margin of ocular sclerite on Rhizoecinae . Postoccipital ridge absent. Ventral midcranial ridge poorly developed or absent. Pronotal ridge and pronotal sclerite absent. Propleural ridge short, not nearly reaching head. Ostioles absent. Glandular pouches absent, although 1 or more long setae may be present in this position on abdominal segment VIII, without associated loculate pores............................................................. 8

s8. Body cylindrical or laterally flattened, somewhat pointed at both ends. Without caudal extensions (anal lobes). Two pairs of simple eyes generally present. Preocular ridges generally present although often weak. Base of antennae placed close together near anterior margin of head. Tarsi generally clearly 2 segmented. Penial sheath clearly extending posteriorly from posterior margin of abdominal segment VIII, often about as broad as long............................ RHIZOECIDAE ( Fig. 5 )

- Body generally showing some modification for transport by ants, either with an enlarged cephalothorax and narrow abdomen or with body laterally flattened. Without simple eyes. Preocular ridges absent. Base of antennae may be either close together or far apart near anterior margin of head. Tarsi 1 segmented. Penial sheath generally lying beneath abdominal segment VIII (except Eumyrmococcus taylori Williams )........................................... XENOCOCCIDAE Tang ( Figs 6 , 7 )

Figures

FIGURE 2. Rhizoecus coffeae Laing. Macropterous male. (Rhizoecidae, Rhizoecina). Where A=hair-like seta, B=fleshy seta, C=loculate pores, K=proximal end of metathoracic leg, and Z=mouth tubercle. For abbreviations, see p. 8.

FIGURE 3. Ripersiella hibisci (Kawai & Takagi). Macropterous male. (Rhizoecidae, Ripersiellina). Where A=hair-like seta, B=fleshy seta, C =loculate pores, F=alar sensoria, and K=proximal end of metathoracic leg. For abbreviations, see p. 8.

FIGURE 4. Rhizoecus dianthi Green. Brachypterous male. (Rhizoecidae, Rhizoecina). Where A=hair-like seta, B=fleshy seta, C=loculate pore, D=simple pore, and G=wing microtrichia. For abbreviations, see p. 8.

FIGURE 5. Kissrhizoecus hungaricus Kozár & Konczné Benedicty. Apterous male. (Rhizoecidae, Rhizoecina). WhereA=hairlike seta, B=fleshy seta, C=loculate pore and K=tibio and tarsus. For abbreviations, see p. 8.

FIGURE 6. Neochavesia nr. trinadadensis (Beardsley). Apterous adult male. (Xenococcidae). H=apex on antenna, K=tibia and tarsus of metathoracic leg, and N=ventral view of penial sheath. For abbreviations, see p. 8.

FIGURE 7. Xenococcus acropygae Williams. Apterous adult male. (Xenococcidae). Where A=minute setae, K=apex of limbs, M=lobes of abdominal segment VIII and Q=antenna. For abbreviations, see p. 8. (Modified after Williams 1998).

FIGURE 8. Phenacoccus solenopsis Tinsley. Macropterous male. (Pseudococcidae, Phenacoccinae). Where C=loculate pores, D=simple pore, J=dorsal view of penial sheath, K=apex of leg, and L=ventral view of penial sheath. For abbreviations, see p. 8.

FIGURE 9. Phenacoccus sp. Evia, Greece. Macropterous male. (Pseudococcidae, Phenacoccinae). Where C=loculate pore and D=simple pore.

FIGURE 10. Paracoccus glaucus (Maskell). Macropterous male. (Pseudococcidae, Pseudococcinae). Where C=loculate pores, D=simple pore, J=dorsal view of penial sheath, K1=tibio-tarsal articulation, and K2=proximal end of metathoracic leg, L=ventral view of penial sheath and N=side view of penial sheath. For abbreviations, see p. 8.

FIGURE 11. Spilococcus atriplicis (Cockerell) Macropterous male. (Pseudococcidae, Pseudococcinae). Where C=loculate pore, D=fleshy seta and E=hair-like seta.

FIGURE 12. Trabutina elastica (Marchal). Macropterous male (Pseudococcidae, Pseudococcinae). Where C=loculate pore, D=fleshy seta and E=hair-like seta.

FIGURE 13. Oracella acuta (Lobdell).Apterous male (Pseudococcidae, Pseudococcinae). Where A=dorsal view of thorax and B=ventral view of thorax, C=loculate pore and D=fleshy seta.

FIGURE 14. Promyrmococcus dilli Williams. Apterous adult male. (Pseudococcidae, Pseudococcinae, Allomyrmococcini). Where C=loculate pore, D=simple pore, J=dorsal view of penial sheath, K=proximal end of metathoracic leg, L=ventral view of penial sheath, P=anterior and posterior views of trochanter, Q=apex of antenna, R=circuli and T=ventral view of right side of head. For abbreviations, see p. 8. (Modified after Williams 2002).

FIGURE 15. Acanthococcus adenostomae (Ehrhorn). Macropterous male. (Eriococcidae, Acanthococcid group). Where A=cranial apophysis, D=fleshy setae and X=pores on head.

FIGURE 16. Acanthococcus melnikensis Hodgson & Trencheva. Macropterous male. (Eriococcidae, Acanthococcid group). Where A=cranial apophysis, D=fleshy seta, E=hair-like seta, M=sclerotised structures associated with genitalia and O=loculate pores.

FIGURE 17. Eriochiton armatus Brittin. Macropterous male. (Eriococcidae, Acanthococcid group). Where A=cranial apophysis and C=loculate pore.

FIGURE 18. Ovaticoccus adoxus (Ferris). Apterous adult male. (Eriococcidae, Acanthococcid group). Where C=loculate pore, D=fleshy seta and F=metacoxa and trochanter.

FIGURE 19. Apiomorpha munita tereticornuta Gullan. Macropterous male. (Eriococcidae, Acanthococcid group). Where A=cranial apophysis, C=loculate pore, D=fleshy seta, E=hair-like seta, N=side view of simple eye, R=dermal reticulation on ocular sclerite and X=pores on head.

FIGURE 20. Apiomorpha ovicola (Froggatt). Macropterous male. (Eriococcidae,Acanthococcid group). WhereA=cranial apophysis, C=loculate pore, D=fleshy seta, E=hair-like seta, N=side view of simple eye and X=pores on head.

FIGURE 21. Apiomorpha pharetrata (Schrader). Macropterous male. (Eriococcidae, Acanthococcid group). Where A=cranial apophysis, D1=fleshy seta on antennae, D2=fleshy seta on tarsus, E=hair-like seta and R=dermal reticulation on ocular sclerite.

FIGURE 22. Apiomorpha rosaeformis (Froggatt). Macropterous male. (Eriococcidae,Acanthococcid group). Where D=fleshy seta and E == hair-like seta.

FIGURE 23. Apiomorpha spinifer Froggatt). Macropterous male. (Eriococcidae, Acanthococcid group). Where A=cranial apophysis, D=fleshy seta, E=hair-like seta, N=side view of simple eye, R=dermal reticulation on ocular sclerite.

FIGURE 24. Dactylopius coccus (Costa). Macropterous male. (Dactylopiidae). Where P=spur-like seta and T=microtrichia.

FIGURE 25. Dactylopius tomentosus (Lamark). Macropterous male. (Dactylopiidae). Where A=cranial apophysis, D=fleshy seta and X=pore-like structure on head.

FIGURE 26. Callococcus leptospermi (Maskell). Macropterous male. (Eriococcidae, Gondwanan group). Where A=cranial apophysis and D=fleshy seta.

FIGURE 27. Calycicoccus merwei Brain. Macropterous male. (Eriococcidae, Gondwanan group). For abbreviations, see p. 8.

FIGURE 28. Capulinia sallei Signoret. Macropterous male. (Eriococcidae, Gondwanan group). Where E=hair-like setae. For abbreviations, see p. 8.

FIGURE 29. Carpochloroides viridis Cockerell. Macropterous male. (Eriococcidae, Gondwanan group). Where C=fleshy seta, D=hair-like seta and K=tibia and tarsus of metathoracic leg. For abbreviations, see p. 8.

FIGURE 30. Choneochiton casuarinae Hodgson, Mille & Cazères. Macropterous male. (Eriococcidae, Gondwanan group). Where A=hair-like setae, B=fleshy seta, C=proximal end of metathoracic leg, D=dorsal view of penial sheath, E=ventral view of penial sheath, and F=ventral view of anterior end of head.

FIGURE 31. Cystococcus echiniformis Fuller. Macropterous male. (Eriococcidae, Gondwanan group). Where A=bifurcated fleshy setae on antenna, D=fleshy setae on leg and abdomen, E=hair-like seta on antennae, G=dermal markings, N=side view of simple eye, and iii–viii=abdominal segmentation,

FIGURE 32. Dromedaricoccus hansoni Hodgson & Miller. Macropterous male. (Eriococcidae, Gondwanan group). Where A=cranial apophysis, B=side view of aedeagus, C=hair-like seta, D=fleshy seta and F=tibia and tarsus of metathoracic leg. For abbreviations, see p. 8.

FIGURE 33. Eriogallococcus isaias Hodgson & Magalhães. Macropterous male. (Eriococcidae, Gondwanan group). Where A=cranial apophysis, C=hair-like seta, D=fleshy seta, E=apex of style and F=tibia and tarsus of metathoracic leg. For abbreviations, see p. 8.

FIGURE 34. Lachnodius?eucalypti (Maskell). Macropterous male. (Eriococcidae, Gondwanan group). Where A=cranial apophysis.

FIGURE 35. Lobimargo williamsi Hardy & Gullan. Macropterous adult male (Eriococcidae, Gondwanan group) (after Hardy et al. 2011). For abbreviations, see p. 8.

FIGURE 37. Pseudotectococcus rolliniae Hodgson & Gonçalves. Macropterous male. (Eriococcidae, Gondwanan group).Where A=cranial apophysis, B=dorsal and ventral view of posterior abdominal segments and penial sheath, C=simple pore, D=fleshy seta, E=apical four segments of antenna and F=tibia and tarsus of metathoracic leg. For abbreviations, see p. 8.

FIGURE 38. Stibococcus cerinus Miller & González. Macropterous male. (Eriococcidae, Gondwanan group) (after Miller & Gonzalez, 1975). Where C=loculate pores of glandular pouch and X=pores on head.

FIGURE 39. Tanyscelis verrucula (Froggatt). Macropterous male. (Eriococcidae, Gondwanan group). Where A=cranial apophysis and X=pores on head.

FIGURE 40. Tectococcus ovatus Hempel. Macropterous male. (Eriococcidae, Gondwanan group). Where X=pores on head.

FIGURE 41. Pseudochermes fraxini (Kalt.). Apterous adult male.(Cryptococcidae; Gondwanan group). (AfterAfifi 1968). Where C=apical antennal segments, F=metathoracic leg and H=side view of posterior abdominal segments and penial sheath. For abbreviations see p. 8.

FIGURE 42. Eriococcus buxi (Fonscolombe). Macropterous male. (Eriococcidae, BSE group) (after Afifi 1968). Where B=anterior view of head, C=apical antennal segments, D=mesoprephragma (anterior view) and mesopostphragma (posterior view), F=metathoracic leg and G=claw.

FIGURE 43. Cylindrococcus spiniferus Maskell. Macropterous male. (Eriococcidae, BSE clade) (after Gullan 1978). Where B=side view of penial sheath. For abbreviations, see p. 8.

FIGURE 44. Mangalorea hopeae Takagi. Macropterous male. (Beesoniidae). (After Raman & Takagi 1992).

FIGURE 45. Gallacoccus secundus Takagi. Macropterous male. (Beesoniidae) (After Takagi 2001). Where B and C=apical antennal segment, D=dorsal and ventral view of abdomen, E=side view penial sheath and F=fleshy setae.

FIGURE 46. Beesonia dipterocarpi Green. Macropterous male. (Beesoniidae). Where D=normal fleshy seta, E=hair-like seta, G=bifurcated fleshy setae on antennae, M=spiracle, P=tarsal campaniform sensilla, X=end of style, ventral view, and Y=end of style, side view.

FIGURE 47. Parastictococcus brachystegiae (Hall). Macropterous male. (Stictococcidae).

FIGURE 48. Parastictococcus hargreavesi (Vayssière). Macropterous male. (Stictococcidae). Where A=cranial apophysis.

FIGURE 49. Parastictococcus multispinosus (Newstead). Macropterous male. (Stictococcidae). Where D=fleshy setae and E=hairlike seta.

FIGURE 50. Parastictococcus?multispinosus (Newstead). Macropterous male. (Stictococcidae). Where A=cranial apophysis.

FIGURE 51. Stictococcus intermedius Newstead. Macropterous male. (Stictococcidae).WhereA=cranial apophysis and L=loculate pore.

FIGURE 52. Stictococcus vayssierei Richard. Macropterous male. (Stictococcidae). Where L=loculate pore. Where A=cranial apophysis, E=divided hair-like seta and L=loculate pore.

FIGURE 53. Conchaspis angraeci Cockerell. Macropterous male. (Conchaspididae). Where F=bifurcated seta, J=proepisternum + cervical sclerite and R=dermal reticulations on ocular sclerite.

FIGURE 54. Conchaspis capensis Linnaeus. Macropterous male. (Conchaspididae). Where P=spur-like seta on tarsus, R=dermal marking on ocular sclerite and T=dermal marking on abdominal tergites and sternites.

FIGURE 55. Conchaspis socialis Green. Macropterous male. (Conchaspididae). Where F=bifurcated seta and R=dermal marking on ocular sclerite.

FIGURE 56.Conchaspis vayssiereiMamet.Macropterous male. (Conchaspididae).Where D=fleshy setae on antenna, F=bifurcated seta, R=dermal markings on ocular sclerite and T=dermal markings on thorax, legs and antennae.

FIGURE 57. Phoenicococcus marlatti Cockerell. Apterous adult male. (Phoenicococcidae).

FIGURE 58. Halimococcus lampas Cockerell. Apterous adult male. (Halimococcidae). (After Stickney 1934). Where B=apex to antenna and C=dorsal view of metathoracic leg.

FIGURE 59. Fiorinia externa Ferris. Macropterous male. (Diaspididae, Diaspidinae, Fioriniina) (After Lambdin et al. 2007). Where A=dermal reticulations on prescutum, B=dermal markings on scutum, C=dermal markings on haltere and dorsum of abdomen, D=side view of penial sheath, E=dermal reticulations on ocular sclerite, F=bifurcated seta, G=distal end of metathoracic leg, H=apical segment of antenna, J=ridges on profemur and K=mouth area. For abbreviations, see p. 8.

FIGURE 60. Leucaspis gigas (Maskell). Macropterous male. (Diaspididae, Aspidiotinae, Leucaspidini). Where C=capitate seta on apex of antenna and F=bifurcated seta. For abbreviation, see p. 8.

FIGURE 61. Labidaspis myersi (Green). Macropterous male. (Diaspididae, Aspidiotini, Leucaspidini). Where F=bifurcated seta.

FIGURE 62. Allokermes galliformis (Riley). Macropterous male. (Kermesidae). Where A=cranial apophysis.

FIGURE 63. Kermes shastensis Ehrhorn. Macropterous male. (Kermesidae). Where A=cranial apophysis, D=fleshy seta and E=hair-like seta.

FIGURE 64. Kermes sp. California, USA. Macropterous male. (Kermesidae). Where A=cranial apophysis.

FIGURE 65. Tachardina aurantiaca (Cockerell). Macropterous male. (Kerriidae). Where A=cranial apophysis.

FIGURE 66. Tachardiella sp. Tampico, Mexico. Macropterous male. (Kerriidae). Where A=cranial apophysis.

FIGURE 67. Cerococcus artemisiae (Cockerell). Macropterous male. (Cerococcidae). Where A=cranial apophysis and R=dermal reticulations on ocular sclerite.

FIGURE 68. Antecerococcus indicus (Maskell). Macropterous male. (Cerococcidae). Where A=cranial apophysis and R=dermal reticulations on ocular sclerite.

FIGURE 69. Antecerococcus ornatus (Green). Macropterous male. (Cerococcidae). Where R=dermal reticulations on ocular sclerite.

FIGURE 70.Asterodiaspis alba (Takahashi).Macropterous male. (Asterolecaniidae).WhereA=cranial apophysis and R=dermal reticulations on ocular sclerite.

FIGURE 71. Bambusaspis delicata (Green). Macropterous male. (Asterolecaniidae). WhereA=cranial apophysis and R=dermal reticulations on ocular sclerite.

FIGURE 72. Bambusaspis longa (Green). Macropterous male. (Asterolecaniidae). Where D=fleshy seta, E=hair-like seta and R=dermal reticulations on ocular sclerite.

FIGURE 73. Asterolecanium petrophilae (Fuller). Macropterous male. (Asterolecaniidae).

FIGURE 74. Hsuia cheni Borchsenius. Macropterous male. (Asterolecaniidae). Where A=cranial apophysis and R=dermal reticulations on ocular sclerite.

FIGURE 78. Aclerda arundinariae McConnell. Macropterous male. (Aclerdidae, Aclerdinae). Where A=cranial apophysis, P=loculate pore and R=dermal reticulations on ocular sclerite.

FIGURE 79. Aclerda distorta Green. Macropterous male. (Aclerdidae, Aclerdinae). Where A=cranial apophysis, E=loculate pore and R=dermal reticulations on ocular sclerite.

FIGURE 80. Aclerda tillandsiae Howell. Macropterous male. (Aclerdidae, Aclerdinae).

FIGURE 81. Aclerda tokionis Cockerell. Macropterous male. (Aclerdidae, Aclerdinae). Where A=cranial apophysis.

FIGURE 82. Aclerda sp. A. Michoacán, Mexico. Macropterous male. (Aclerdidae, Aclerdinae). Where A=cranial apophysis, C=loculate pore, D=simple pore and R=dermal reticulations on ocular sclerite.

FIGURE 83. Aclerda sp. B. South Dakota, USA. Macropterous male. (Aclerdidae, Aclerdinae). Where A=cranial apophysis, C=loculate pore, D=simple pore and R=dermal reticulations on ocular sclerite.

FIGURE 84. Lecanaclerda macropoda Hodgson & Millar. Macropterous male. (Aclerdidae. Aclerdinae). Where A=cranial apophysis, D=simple pore, K=apex of antennal capitate seta and R=dermal reticulations on ocular sclerite.

FIGURE 85. Nipponaclerda biwakoensis (Kuwana). Apterous male. (Aclerdidae, Aclerdina) (after McConnell, 1954). Where H=antenna, I=anterior leg, J=posterior spiracle, K=metathoracic leg, and L=ventral view of penial sheath.

FIGURE 86. Rhodesaclerda insleyae Hodgson & Millar. Macropterous male. (Aclerdidae, Rhodesaclerdinae).Where D=simple pore and R=dermal reticulations on ocular sclerite.

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.