ERIOCOCCIDAE

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 : 72-74

Group B. GONDWANAN ERIOCOCCIDAE

Introduction. This group includes a wide variety of genera all of which lack hamulohalteres and most lack alar setae (present on Choneochiton ) but do have postmesospiracular setae. All have the campaniform sensilla on each trochanter more or less in a line. Almost all of these genera are gall-inducing and have various structural modifications to allow access to the female for mating, such as very elongate abdomens. The genera considered to belong to this group for which the males are known are: Callococcus Ferris ; Calycicoccus Brain ; Capulinia Signoret ; Carpochloroides Cockerell ; Choneochiton Hodgson ; Cystococcus Fuller ; Dromedaricoccus Hodgson & Miller ; Eriogallococcus Hodgson & Magalhãaes ; Lachnodius Maskell ; Tanyscelis Hardy & Gullan ; Pseudomontanococcus Kozár & Hodgson ; Pseudotectococcus Hempel ; Stibococcus Miller & González and Tectococcus Hempel. Of these, Calycicoccus merwei Brain is the only indigenous eriococcid known from Africa, Choneochiton is restricted to New Caledonia, and Capulinia , Dromedaricoccus , Eriogallococcus , Pseudotectococcus and Tectococcus are all from South America; the remainder are restricted to Australia.A key for separation of the known males of South American genera can be found in Hodgson and Miller (2010). A key to the adult males of all of the genera in this group is provided below.

Group diagnosis based on macropterous adult male morphology of Callococcus , Calycicoccus , Capulinia , Carpochloroides , Choneochiton , Cystococcus , Dromedaricoccus , Eriogallococcus , Lachnodius, Lobimargo , Pseudotectococcus , Stibococcus , Tanyscelis and Tectococcus species. Body often attenuated with abdomen generally narrowing posteriorly; body setae generally all hs, occasionally some stoutly fs ( Callococcus, Lobimargo and Pseudotectococcus ); loculate pores only present in glandular pouches. Head: simple pores sometimes present on head; ocular sclerite with few or no reticulations; simple eyes often quite large; preocular ridge well developed, with both anterior and posterior arms; interocular ridge absent; genal setae usually present (absent on Lachnodius ); postoccipital ridge well developed, usually with both anterior and posterior arms; number of antennal segments often reduced, ranging from 4–10 segments; capitate setae generally present on antennal segments in addition to those on apical segment (exceptions Callococcus, Lobimargo and Tectococcus ); flagellar segments with both hs and fs, latter generally barely differentiated from hs but quite long. Thorax: prescutum usually with prescutal setae; scutal setae present laterally; scutellum with scutellar setae; postemesospiracular setae present and often abundant (few in Cystococcus ); metasternum usually with few setae; metaprecoxal ridge absent; postmetaspiracular setae usually present, often abundant; hamulohalteres absent; alar setae absent (present on Choneochiton and Tectococcus ); alar sensoria absent; trochanter with campaniform sensoria in a line on each side; fs present or absent on legs; tibia with 2+ spurs; tarsi mainly 2 segmented; claw digitules capitate; claw with a denticle. Abdomen: tergites and sternites usually poorly sclerotised or unsclerotised; fs absent or almost indistinguishable from hs (except Callococcus, Lobimargo and Pseudotectococcus ); caudal extensions occasionally present; glandular pouches on segment VIII present or absent; abdominal segment IX distinct but fused to style; anus usually distinct in centre of segment IX; penial sheath with most setae quite long; style becoming sharply pointed apically and generally about twice as long as basal width.

In addition to the genera diagnosed above, Pseudomontanococcus martini Kozár & Konczné Benedicty also belongs to this group but is apterous.

Key to some “Gondwanan” Eriococcidae based on adult male morphology.

1. Apterous. Glandular pouches absent. Body setae numerous and long. Antennae short, 9 segmented, each segment much shorter than wide ( Papua New Guinea).................... Pseudomontanococcus martini Kozár & Konczné Beneticty ( Fig. 36 )

- Macropterous. Other characters not in this combination....................................................... 2

2. Glandular pouches absent. Abdomen greatly attenuated....................................................... 3

- Glandular pouches present. Adomen generally not attenuated.................................................. 6

3. Antennae 4 segmented, densely covered in setae. Fleshy setae on antennae sometimes branched. With 4 or more capitate setae on apical antennal segment. Abdomen exceedingly long, 3 or more times longer than head + thorax (Australasia)................................................................................... Cystococcus Fuller spp. ( Fig. 31 )

- Antennae 8 or 9 segmented. Fleshy setae on antennae not branched. Apical antennal segment with 4 or fewer capitate setae. Abdomen, even if attenuated, less than 2 times length of head + thorax.......................................... 4

4. Antennae 9 segmented. All antennal setae very long, much longer than 2x antennal width. Capitate setae on antennae restricted to apical segment. Tarsi 1 segmented. Postmesopiracular setae abundant (Australasia)................................................................................................... Tanyscelis verrucula (Froggatt) ( Fig. 39 )

- Antennae 8 segmented. Setae on antennae not 2x longer than antennal width. Capitate setae present on some antennal segments other than apical segment. Other characters not in this combination............................................. 5

5. Fleshy setae on antennae broad and fairly short. Pores absent from head. Tarsi 1 segmented. Alar setae absent (Australasia).................................................................. Callococcus leptospermi (Maskell) ( Fig. 26 )

- Fleshy setae similar to hair-like setae. Simple pores frequent on head. Tarsi 2 segmented. Alar setae present (Neotropical)......................................................................... Tectococcus ovatus Hempel ( Fig. 40 )

6. Antennae 5–8 segmented. Tarsi 1 segmented............................................................... 7

- Antennae 10 segmented. Tarsi 1 or 2 segmented........................................................... 10

7. Fleshy setae on antennae short and stout. Fleshy setae present on dorsum and venter of abdomen, each short and stout. Basisternal setae present but few............................................................................. 8

- Fleshy setae basically similar to hair-like setae and hard to differentiate. Fleshy setae absent from abdomen or, if considered to be present, not short and stout. Basisternal setae abundant..................................................... 9

8. Genal setae setose and few. Postmesospiracular setae few and hs ( South Africa)....... Calycicoccus merwei Brain ( Fig. 27 )

- Genal setae fleshy and abundant. Postmesospiracular setae fleshy and abundant (Neotropical)........................................................................................... Pseudotectococcus Hempel spp. ( Fig. 37 )

9. Penial sheath narrow and elongate, almost as long as abdomen and about 4x as long as basal width. Antennae 5 segmented. Prosternal setae few or absent (Neotropical).................... Eriogallococcus isaias Hodgson & Magalhães ( Fig. 33 )

- Penial sheath short and narrow, length about 1.5x longer than basal width. Antennae 6 segmented. Prosternal setae abundant (Neotropical)............................................. Dromedaricoccus hansoni Hodgson & Miller ( Fig. 32 ) -

10. Penial sheath about 3x longer than basal width. Tarsi 2 segmented. Claw without a denticle. Antennal setae long, about 2x longer than antennal width (Neotropical)....................................... Capulinia sallei Signoret ( Fig. 28 )

- Penial sheath much shorter than 3x basal width. Other characters not in this combination........................... 11

11. Genal setae absent. Ocelli positioned away from postocular ridge, in centre of ocular sclerite (Australasia)...............

.................................................................. Lachnodius ?eucaplyti (Maskell) ( Fig. 34 ) - Genal setae abundant. Ocelli positioned close to or touching postocular ridge.................................... 12

12. Antemesospiracular setae abundant. Body setae either capitate or with a blunt apex. Capitate setae on antennae present on nonapical segments (Oceania)............................. Choneochiton casuarinae Hodgson, Mille & Cazères ( Fig. 30 )

- Antemesospiracular setae absent. Setae fleshy or hair-like, neither capitate nor with a blunt apex. Capitate setae on antennae, if present, restricted to apical segment (or if considered present, hard to differentiate from other setae).................. 13

13. Glandular pouch setae long, extending posteriorly past apex of penial sheath. Dorsal abdominal setae more abundant than ventral abdominal setae. Small pores absent from head. Loculate pores of glandular pouches restricted to within each pouch (Neotropical)...................................................... Carpochloroides viridis Cockerell ( Fig. 29 )

- Glandular pouch setae short, only extending to about half length of penial sheath. Dorsal abdominal setae not as abundant as ventral abdominal setae. Small pores present anteriorly on head. Loculate pores of glandular pouches extending out onto derm on either side of pouch............................................................................... 14

14. Fleshy setae on antennae long, more than twice width of antennal segments. Metasternum with frequent setae. Caudal extensions on abdomen absent. Tergites and sternites of abdominal segment VIII unsclerotised (Australasian)............................................................................... Lobimargo williamsi Hardy & Gullan ( Fig. 35 )

- Fleshy setae on antennae absent, setae on antennae all hs and short, length subequal to width of antennal segments. Metasternum without metasternal setae. Caudal extensions on abdomen present and rounded. Tergites and sternites of abdominal segment VIII present (Neotropical).................................... Stibococcus cerinus Miller & González ( Fig. 38 )

Figures

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 36. Pseudomontanococcus martini Kozár & Konczné Benedicty. Apterous adult male. (Eriococcidae, Gondwanan group). Where A=cranial apophysis, D=fleshy setae, E=hair-like setae, R=dermal reticulations on ocular sclerite and X=pores on head.

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.