NEOCOCCOID

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

3. Legs well developed but tibia + tarsus fused. Antennae 3 segmented. Each caudal extension on abdominal segment VIII sclerotised, forming an oval process. Bilocular pores present. Penial sheath much longer than basal width........................................................................................ MICROCOCCIDAE Leonardi (Fig. 77)

- 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)

DESCRIPTIONS AND DIAGNOSES OF NEOCOCCOID TAXA BASED ON ADULT MALE MOR- PHOLOGY

RHIZOECIDAE Williams, 1969

Rhizoecini Williams 1969, 335.

Rhizoecinae Williams 1969, 335; Koteja 1974b, 45; Tang 1992, 42.

Rhizoecidae Williams; Hodgson, 2012, 4.

Type genus: Rhizoecus Künkel d’Herculais.

Type species: Rhizoecus falcifer Künkel d’Herculais 1878, 164 by monotypy.

Introduction. The taxa with the hypogaeic and myrmecophilous mealybugs have, until recently, been included in the Pseudococcidae as the subfamily Rhizoecinae. However, a phylogenetic study based on DNA sequences and including representatives from all five pseudococcid subfamilies, found that the Rhizoecinae (represented by 4 species in 3 genera) formed a monophyletic group sister to the remaining pseudococcids ( Downie & Gullan 2004). Later, based on adult male characters only, Hodgson and Foldi (2005) found that Rhizoecus formed a separate clade with Pityococcus, sister to the Pseudococcidae. Since then, also largely based on the morphology of the adult males, Hodgson (2012) reviewed the relationships of the Pseudococcidae and the hypogaeic and myrmecophilous mealybugs, and concluded that the rhizoecine mealybugs formed a separate family from the Pseudococcidae ( Rhizoecidae Williams) and that this family included two subfamilies, Rhizoecinae Williams and Xenococcinae Tang. A more recent study ( Hodgson & Hardy 2013), also based on adult male morphology, also found the Rhizoecidae to be a separate clade, sister to the Pseudococcidae. However, Danzig and Gavrilov-Zimin (2014) did not accept this and argued that: (a) the hypogaeic mealybugs belonged within the Pseudococcidae, possibly sister to Mirococcopsis Borchesenius, and that (b) the myrmecophilous Xenococcinae were unrelated to the Pseudococcidae and might be closest to the Margarodidae s.l. Danzig and Gavrilov-Zimin therefore raised the Xenococcinae to family status, Xenococcidae Tang, although this was without phylogenetic support. Whilst it is still the view of the present author that the Rhizoecidae are sister to the Pseudococcidae, he agrees that the relationships of the Xenococcinae are problematic and that it should be raised to family status. As understood here, the Rhizoecidae contains 16 genera and 214 species ( García Morales et al. 2019).

Family diagnosis based on adult male morphology. Many species apterous but some with brachypterous or fully macropterous adult males. Unless otherwise stated, diagnosis refers to all morphs. Body generally small to minute, cylindrical, somewhat pointed at both ends. Head without ocelli, although two pairs of simple eyes generally present; preocular ridges generally recognisable ventrally although often not strongly developed; postocular ridge absent although a faint line occasionally present marking posterior margin of ocular sclerite; postoccipital ridge absent; dorsal midcranial ridge absent; ventral midcranial ridge poorly developed or absent; base of antennae placed close together near anterior margin of head; number of antennal segments variable, each 3–10 segmented, with fs and hs setae on most segments; capitate setae on antennae generally absent or not differentiated; neck entirely absent. Loculate pores frequently present on head, thorax and/or abdomen. Thorax. Pronotal ridge and pronotal sclerite absent; propleural ridge short, at most only extending anteriorly about half-way to head; apterous species without any mesothoracic sclerotisation. Metaprecoxal ridge absent; leg setae hs or spinose, not fs; each trochanter with 3 round sensoria arranged in a curve or triangle on each side; trochanter often divided by a Y-shaped sclerotisation; tarsi with 2 spurs; tarsi generally clearly 2 segmented, occasionally 1 segmented; tarsal digitules often undifferentiated; claws long and narrow with setose digitule s. Abdomen. Ostioles absent; abdominal segment VIII without glandular pouches and without a group of disc-pores, although 1 or more long setae may be present in this position; abdominal segment IX not apparently differentiated, but position indicated by anal opening found dorsally at anterior end of dorsal ridge on penial sheath; ventral opening of penial sheath often indistinct and never with lateral finger-like processes; penial sheath about as broad as long; penial sheath without a large, U-shaped, sclerotised structure extending anteriorly within abdomen from base of aedeagus (if an obvious internal sclerotisation present, this never as long as penial sheath). Macropterous and brachypterous species also with: scutum without a membranous area; basisternum without a median ridge; alar lobes present on each wing but hamulohalteres absent; alar setae present; alar sensoria present or absent.

Discussion. In his 2012 paper, Hodgson found that the morphology of the adult males of the Rhizoecidae was very variable, particularly in the number of antennal segments, and that male morphology did not entirely support the classification based on the morphology of the adult female of Kozár & Konczné Benedicty (2007). The males of the macropterous rhizoecine mealybugs, Rhizoecus coffeae Laing ( Fig. 2), Ripersiella hibisci ( Figs 2 & 3) and Rhizoecus dianthae Green ( Fig. 4), plus an apterous rhizoecine mealybug, Kissrhizoecus hungaricus Kozár & Konczné Benedicty ( Fig. 5), are illustrated here to illustrate the diversity of the family. A key to the described rhizoecine adult males plus descriptions and other illustrations can be found in Hodgson (2012).

XENOCOCCIDAE Tang 1992

Xenococcinae Tang 1992, 42.

Xenococcidae Tang; Danzig & Gavrilov-Zimin 2014, 45. Revised status.

Type genus: Xenococcus Silvestri 1924, 312.

Type species: Xenococcus annandalei Silvestri by monotypy and original designation.

Introduction. The Rhizoecinae ( Rhizoecidae) was divided into two tribes by Tang (1992) on the basis of the presence (Rhizoecini—here Rhizoecidae) or absence (Xenococcini—here Xenococcidae) of ostioles on the adult females. However, it is known now that some adult females of Rhizoecidae lack ostioles (e.g. Capitisetella and Pseudorhizoecus). Nonetheless, in the Rhizoecidae, the third-instar female is a normal feeding instar whereas it has been shown ( Williams 2004) that all three genera included in the Xenococcinae have a third-instar pupal (non-feeding) stage before the fourth-instar (adult) female stage, clearly separating the two subfamilies. In addition, Williams (2004) showed that, in those species where circuli were present, they are flat or bulbous distally in the Rhizoecinae, whereas in the Xenococcinae they are depressed or cup-shaped at the centre of the distal end. Adult female Xenococcinae also lack tubular ducts, pores and tubular cerores, all of which occur on most Rhizoecinae. More recently, Danzig and Gavrilov-Zimin (2014) have reviewed the above classification and concluded that the myrmecophilous mealybugs are unrelated to the “true” mealybugs and are closer to the Margarodidae s.l. due to the presence of the pupal stage in the female life cycle and the absence of tubular ducts, pores and tubular cerores. They therefore raised this group to family level, Xenococcidae Tang, suggesting that it had a common ancestor within the Margarodidae s.l., i.e., outside the neococcids, although this was not supported by phylogenetic evidence. However, Downie and Gullan (2004) included Neochavesia caldasiae (Balachowsky) in their phylogenetic analysis based on molecular studies, and recovered it within the Rhizoecinae, sister to the remaining three hypogaeic species. It is here considered that the Xenococcidae are neococcids and that they probably arose from a common ancestor close to the true mealybugs. However, the final relationships of this family may only be discovered using a more comprehensive phylogenetic study using molecular data and a larger sample of taxa. Currently, the Xenococcidae contain 3 genera ( Eumyrmococcus Silvestri 1926, Neochavesia Williams & Granara de Willink 1992 and Xenococcus Silvestri 1924) and 38 species, although it seems likely that the family is non-monophyletic.

Family diagnosis based on adult male morphology. Entirely apterous and therefore never with any sclerotised plates on mesothorax; body generally showing some modification for transport by ants, either rather narrow or with an enlarged cephalothorax and narrow abdomen. Head without both simple eyes and ocelli; antennae 1–5 segmented; pedicel perhaps always absent; antennae with hs and bristles; preocular ridges absent; postocular ridge absent; postoccipital ridge absent; dorsal midcranial ridge absent; ventral midcranial ridge poorly developed or absent; base of antennae may be either close together or far apart near anterior margin of head; neck constriction absent. Thorax. Barely differentiated from rest of body; pronotal ridge and pronotal sclerite absent; propleural ridge short, at most only extending anteriorly about half-way to head; loculate pores generally absent; legs generally short, often with some segments fused; trochanter with 2 round sensoria on each side; leg setae all hs; trochanters without a strong Yshaped sclerotised ridge; tarsi 1 segmented; tarsal digitules either undifferentiated or setose; claws without a denticle; claw digitules either absent or not capitate. Abdomen. Ostioles absent; abdominal segment VIII without glandular pouches and without a group of disc-pores, although 1 or more long setae may be present in this position; abdominal segment IX not apparently differentiated; penial sheath significantly longer than broad; penial sheath with a large, U-shaped, sclerotised structure extending within abdomen anteriorly from base of aedeagus, this usually longer than length of penial sheath; aedeagus often longer than penial sheath and sharply pointed.

Comment. Illustrations of two myrmecophilous mealybugs are included, namely Neochavesia nr. trinidadensis (Beardlsey) ( Fig. 6), and Xenococcus acropygae Williams ( Fig. 7). Further descriptions and figures of adult male Xenococcidae can be found in Williams (1998, 2004); Kishimoto-Yamada et al. (2005) and Hodgson (2012) and keys to separate the known adult males of the Xenococcidae are given in Hodgson (2012).

Discussion: Schneider and LaPolla (2011) showed that all Xenococcidae are obligate trophobionts with Acropyga Roger ants. These ants are considered to feed primarily on the honeydew produced by the xenococcine scale insects and the alate ant queen is known to carry one of the xenococcines in its mandibles during her maiden flight, so that the colony she founds has a food source.

The most recent key to the adult females in this (sub)family is in Schneider and LaPolla (2011). In their phylogenetic study of the tribe Xenococcini based on adult female morphology and using both Maximum Parsimony and Bayesian Inference, Schneider and LaPolla (2011) obtained a polytomy with four clades and one ungrouped species, with one clade equating to Neochavesia, one to Xenococcus and three to Eumyrmococcus, which therefore may not be monophyletic. These clades were found to be related to their geographical distributions, with Neochavesia restricted to the Neotropics, Xenococcus to the Indo-Australian Region, the scorpioides- clade (three species, including the ungrouped E. williams Kozár & Konczné Benedicty) of Eumyrmococcus to Africa and the Palaearctic, and the smithii -clade to Asia, the Pacific and Australia. However, Schneider and LaPolla (2011) only used three representative species of pseudococcine and rhizoecine genera ( Geococcus, Rhizoecus and Phenacoccus) as their outgroups. There was support for three of the clades with Bayesian posterior probability scores of 100, although Bootstrap values in the Maximum Parsimony tree were all low (≤76) and there was no support for E. williamsi grouping with the other two species of the E. scorpioides -group. All known adult male Xenococcinae are apterous but the adult males of the three genera are rather different morphologically.

Key to genera of Xenococcidae based on adult male morphology:

1. Antennae 1 segmented. Legs reduced, each with a spine-like pointed claw and two spine-like digitules. Lobes on abdominal segment VIII small, each with a few short setae + 3 longer setae. Head, thorax and 1st-abdominal segments all showing some strong sclerotisation................................................... Xenococcus acropygae Williams ( Fig. 7)

- Antennae 2 or 5 segmented. Legs generally well developed but occasionally reduced, but claw digitules (when present) setose and structure of claw variable but not spine-like. Lobes on abdominal segment VIII either absent or well developed, without the above combination of setae. Without any strong sclerotisation on head, thorax or 1^st- abdominal segment................ 2

2. Abdominal segment VIII with well-developed lateral finger-like processes on either side of ventral opening of penial sheath, each lobe with many setae of moderate-length. Abdomen with a strong constriction between segments II and III. Claw digitules present, setose.............................................. Neochavesia Williams & Granara de Willink ( Fig. 6)

- Abdominal segment VIII without lateral finger-like processes on either side of ventral opening of penial sheath. Setae on abdominal segment VIII either very few and extremely minute or with at least a few setae longer than penial sheath. Abdomen more or less tapering posteriorly, without a strong constriction between segments II & III. Claw digitules absent......................................................................................... Eumyrmococcus Silvestri

PSEUDOCOCCIDAE Cockerell 1905

Pseudococcini Westwood; Cockerell 1905, 193.

Pseudococcinae Westwood; Leonardi 1920, 375.

Pseudococcidae Westwood; Lobdell 1930, 209.

Type genus: Pseudococcus Westwood 1840.

Type species: Dactylopius longispinus Targioni Tozzetti 1867.

Introduction. The Pseudococcidae is the second largest family within the Coccomorpha, with 259 genera and 1989 species ( García Morales et al. 2019). It includes two subfamilies, the Phenacoccinae and the Pseudococcinae ( Williams & Gullan 2010) [note that the hypogaeic and myrmecophilous mealybugs are here considered to be separate families, Rhizoecidae and Xenococcidae. For keys to separate the Rhizoecidae from the Pseudococcidae, see Key on p. 20. The Phenacococcinae contain about 50 genera and the Pseudococcinae about 200 genera ( Hardy et al. 2008). Currently, there is no satisfactory or generally accepted suprageneric classification for the mealybugs ( Downie & Gullan 2004; Hardy et al. 2008; Kaydan et al. 2015; Danzig & Gavrilov-Zimin 2014). Williams and Gullan (2010) listed 4 family-group names belonging to the Phenacoccinae (excluding the Rhizoecidae) and 12 belonging to the Pseudococcinae. Danzig and Gavrilov-Zimin (2014) divide the Phenacoccinae into 8 generic groups (but they included Puto Signoret in the Pseudococcidae, here considered to belong to the Phenacoleachiidae), and divide the Pseudococcinae into 11 generic groups, one of which is the Rhizoecus group (here considered a separate family). It is likely that all suprageneric classifications will undergo significant changes in the future. The most constant such groups within the Pseudococcinae are the Trabutinini, Planococcini and Pseudococcini. In addition to these tribes, Williams (1978) introduced the tribe Allomyrmococcini, within the Pseudococcinae, to take various genera of anomalous ant-attended mealybugs. Although immature male stages are known for a few species of Allomyrmococcini ( Williams 1978), the only adult male known in this tribe is that of Promyrmococcus dilli Williams ( Williams 2002; Hodgson 2012), which is apterous ( Fig. 14)

Generic diagnosis based on adult male morphology. Most species probably alate but some known to be brachypterous and/or apterous; body generally small. Unless otherwise stated, diagnosis refers to all morphs. Head with ocelli in addition to two pairs of simple eyes; postocular ridge generally distinct and present even on apterous species; postoccipital ridge present or absent; dorsal and ventral midcranial ridges usually well developed; ocular sclerite with striations or concentric ridges restricted to around simple eyes; number of antennal segments variable but each antenna usually 9 or 10 segmented, with fs and hs setae on most segments; capitate setae on antennae generally absent or poorly differentiated; neck constriction present or absent. Thorax. Pronotal ridge and pronotal sclerite generally present; prescutum with prescutal setae; postmesospiracular setae generally present; loculate pores frequently present on thorax and/or abdomen; hs and fs setae on legs hard to differentiate; each trochanter with 3 round sensoria on each side in a curve or triangle; trochanter often divided by a Y-shaped sclerotisation; tarsi generally clearly 2 segmented, occasionally 1 segmented; tarsal digitules capitate; claws long and narrow with setose digitules; tarsi generally with 2 or more spurs. Abdomen. Ostioles often present between abdominal segments VI and VII; glandular pouches usually present on abdominal segment VIII, each with a group of loculate pores; abdominal segment IX present as a sclerite on top of penial sheath; anal opening facing posterodorsally at posterior end of segment IX; ventral opening of penial sheath large and distinct, often with a lobe on each lateral margin; penial sheath usually about as broad as long. Alate species generally also with scutum without a median membranous area; scutum with scutal setae medially and laterally; scutellum transversely rectangular; prosternal ridges absent; metaprecoxal ridge well developed; halteres, alar lobes, alar setae and alar sensoria present on each wing; basisternum without a median ridge.

Comment. Four keys to adult male pseudococcids have been published to date ( Beardsley 1960; Afifi & Kosztarab 1967; Afifi 1968; Hodgson 2005) although the descriptions and illustrations of the males of some Hawaiian species in Beardsley (1960) are incomplete and so the inclusion of them here is tentative. Thus, the following key is a preliminary attempt to divide the pseudococcids into higher groups based entirely on male characters.

Key to known adult male Pseudococcidae

1. Apterous or brachypterous............................................................................. 2 - Macropterous........................................................................................ 7

2. Antennae 8 or 9 segmented............................................................................. 3 - Antennae 10 segmented................................................................................ 6

3. Pronotal ridge present. Antennae 8 segmented. Dorsal sclerites present on thoracic segments. Glandular pouch well developed..................................................... Saccharicoccus sacchari (Cockerell) (apterous form)

- Pronotal ridge absent. Antennae 8 or 9 segmented. Dorsal sclerites absent on thoracic segments. Glandular pouch poorly developed, without a pouch but with glandular pouch setae and a few loculate pores.................................. 4

4. Body with exceptionally long setae plus a fairly dense covering of short setae on all body segments. Abdominal segments posterior to segment VI all strongly sclerotised. Antennae 9 segmented............. Promyrmococcus dilli Williams ( Fig. 14) ( Allomyrmococcini)

- Body setae all short and sparse. Abdominal segment VII and VIII membranous. Antennae 8 or 9 segmented............ 5

5. Antennae 9 (rarely 8) segmented. Postocular ridge absent. Loculate pores absent apart from near each glandular pouch. Denticle present on claw (brachypterous form with small wing-buds and slight sclerotisation of mesothorax).......... Mirococcus ( Polystomophora) ostiaplurimus (Kiritchenko) (apterous and brachypterous forms) §

- Antennae 8 segmented. Postocular ridge present near each ocellus but short. A few loculate pores present on thorax. Denticle absent from claw............................................................... Asaphococcus agninus Cox.

6. Interocular ridge present, ventral to each ocellus. Pronotal ridge present. Ocular sclerite unsclerotised....... Chorizococcus multiporus Gertsson & Hodgson

- Interocular ridge absent. Pronotal ridge absent. Ocular sclerite sclerotised....... Paramococcus venezuelanus Foldi & Cox

7. Lateropleurite broad and triangular. Length of glandular pouch setae short, less than 1/5th total body length. Tarsi with two tarsal spurs only. Claws often with a denticle...................................................................... 8

- Lateropleurite narrow, only present as a narrow sclerite along lateral margin of basisternum. Glandular pouch setae long, more than 1/5th total body length. Tarsi with more than 2 tarsal spurs distally. Claws without a denticle … Pseudococcinae...... 13

8. Wings without either alar setae or alar pores. Loculate disc-pores absent. Lateral ocelli vestigial. Postmesostigmatic setae absent. Ostioles absent. With only a single pair of glandular pouches.......................... Nairobia bifrons De Lotto

- Wings with both alar setae and alar pores. Loculate disc-pores present. Lateral ocelli well-developed. Postmesostigmatic setae present. Ostioles usually present. With two pairs of glandular pouches, one pair on abdominal segment VII and the other on VIII … Phenacoccinae.................................................................................... 9

9. Capitate setae present on apical three antennal segments. Abdominal segment IX with rather spinose sclerotised pleurites...

............................................................................ Phenacoccus dearnessi King - Capitate setae absent from all antennal segments. Abdominal segments without sclerotised pleurites.................. 10

10. Lateral margins of scutellum not forming a posterior notal wing process. Ostioles poorly developed.....................

................................................................................ Ceroputo pilosellae Šulc - Lateral margins of scutellum extending laterally to form part of a posterior notal wing process. Ostioles well developed.. 11

11. Loculate pores absent from venter of abdomen. Hair-like setae present on dorsum of prothorax. Loculate pores rarely present dorsally on head...................................................... Phenacoccus solenopsis Tinsley ( Fig. 8)

- Loculate pores present on venter of abdomen. Hair-like setae absent dorsally on prothorax. Loculate pores present dorsally on head............................................................................................... 12

12. Preocular ridge very short dorsally. Postoccipital ridge poorly developed or absent. Postmesospiracular setae and loculate pores present medially anterior to basisternum.............................................. Pelionella balteata (Green)

- Preocular ridge quite long dorsally, fusing with fairly distinct postoccipital ridge. Postmesospiracular setae absent anterior to basisternum.................................................... Coccidohystrix ( Centrococcus) insolita (Green)

13. Glandular pouches with 5 glandular pouch setae, 2 long, 2 of intermediate length and 1 short. Capitate setae present on antennal segments III–X. Abdominal disc-pores present only on abdominal segment I........................................................................... Octococcus africanus (Brain); Atrococcus groenlandensis Gertsson & Hodgson

- Glandular pouches each with only 2–4 glandular pouch setae, 2 long and 0–2 medium to short. Capitate setae restricted to apical segments or possibly absent. Abdominal disc-pores usually also present on other abdominal segments................. 14

14. Postocular ridge short, not reaching dorsal simple eye. Anterior margin of basisternum absent. Furca short. Fleshy setae on antennae short, about as long as width of segments......................................................... 15

- Postocular ridge reaching dorsal simple eye and usually extending well past it. Prescutal suture present. Anterior margin of basisternum well-developed and sclerotised. Furca long. Fleshy setae on antennae usually much longer than width of each segment.............................................................................................. 16

15. Basisternum about as long as wide. Lateropleurite very narrow. Postmesospiracular setae absent medially anterior to basisternum......................................................... Saccharicoccus sacchari (Cockerell) (alate form)

- Basisternum clearly wider than long. Lateropleurite quite broad. Postmesospiracular setae present medially anterior to basisternum.......................................................................... Brevennia rehi (Lindinger)

16. Fleshy setae present on body as well as on antennae and legs, although sometimes few on head only ( Pseudococcinae)... 17

- Fleshy setae entirely absent from body although present on antennae and sometimes on legs........................ 27

17. Ventral opening to style bordered by a pair of finger-like processes, each distinctly longer than broad................. 18

- Ventral opening to style either not bordered by a pair of finger-like processes or these short, about as long as wide....... 19

18. Antennae 8 or 9 segmented, very short. Dorsal fleshy setae on abdomen short, 12–14 µm long.........................

......................................................................... Palmicultor palmarum (Ehrhorn) - Antennae 10 segmented. Dorsal fleshy setae on abdomen mostly more than 20 µm long.. Pseudococcus spp. ( Ps. antricolens Ferris; P. floriger Ferris; Ps. longispinus (Targioni Tozzetti); Ps. lycopodii Beardsley; Ps. pipturicolus Beardsley) *

19. Fleshy setae common to abundant on both dorsum and venter of abdomen....................................... 20

- Fleshy setae scarce or absent on dorsum and venter of abdomen but present on head............................... 22

20. Setae absent from anterior half of ventral surface of penial sheath, setae only present on finger-like processes ventrally and on

apex of style.................................................................... Clavicoccus tribulus Ferris - Several setae present on anterior half of ventral surface or penial sheath........................................... 21

21. Fleshy setae abundant on gena................................ Ps. comstocki (Kuwana); Ps. cryptus Hempel; Ps. fragilis (Brain); Ps. montanus Ehrhorn; Ps. viburni (Signoret); Chlorococcus ( Ps.) peleae (Beardsley); Dysmicoccus boninsis (Kuwana); D. neobrevipes Beardsley; D. grassii (Leonardi) *

- Fleshy setae few, less than 5 on each gena............................................. Leptococcus metroxyli Reyne

22. Fleshy setae present on abdomen, but very few................................................................23

- Fleshy setae on abdomen absent, fleshy setae only present on head............................................... 24

23. Fleshy setae on antennae short, 18 µm or less...................................... Laminicoccus pandani (Cockerell)

- Fleshy setae on antennae longer, 25 µm or more.... Nesopedronia acanthocauda (Beardsley), N. cibotti (Beardsley), N. crypta (Beardsley), N. hawaiiensis (Ferris); Nesococcus pipturi Ehrhorn *

24. Setae on apex of penial sheath short, shorter than those on body..................................................25

- Setae on apex of penial sheath long, similar to those on abdomen ….............................................. 26

25. Fleshy setae on head and limbs short, much shorter than hair-like setae ….............. Macrocepicoccus loranthi Morrison

- Fleshy setae on head and limbs longer, subequal in length to hair-like setae …......... Plotococcus eugeniae Miller & Denno

26. Antennae quite short, less than 500 µm long. Fleshy setae on antennae about 26 µm long …... Pseudotrionymus refertus Ferris

- Antennae more than 650 µm long. Fleshy setae on antennae about 36 µm long …...................................... ............................................. Chlorococcus straussiae (Ehrhorn); Phyllococcus oahuensis (Ehrhorn) *

27. Fleshy setae on antennae and legs long, about twice as long as antennal width and longer than hair-like setae on body. Dorsal arm of mid-cranial ridge absent. Prosternal pores absent. Abdominal disc pores restricted to margins …........................................................................................... Ferrisia virgata (Cockerell) ( Ferrisia group)

- Fleshy setae on antennae and legs shorter, usually about as long as width of antennal segments and equal to or shorter than hairlike setae on body. Other characters not in this combination … (Trabutinini and Planococcini) …....................... 28

28. Abdomen narrowing significantly about segment III or IV, becoming very narrow ….................................29 - Abdomen of about equal width along most of its length, only narrowing about segment VI or VII ….................... 30

29. Penial sheath very long and narrow, about 10x longer than basal width. Ventral eyes on a protuberance. Most abdominal sternites heavily sclerotised …............................................ Quadrigallicoccus lauracearum Williams & Miller

- Penial sheath short, length shorter than abdominal segment VIII. Ventral eyes not on a protuberance. Abdominal sternites poorly defined …...................................................................? Sphaerococcus tomentosus Fuller

30. Postmesospiracular setae and/or pores present mesad to mesothoracic spiracle, generally extending across entire segment, not restricted to a group posterior to mesothoracic spiracle. Lateral finger-like processes on each side of ventral opening of style well developed and longer than wide.............................................................................. Nipaecoccus bromelicola von Ellenrieder et al. , N. floridensis Beardsley, N. nipae (Maskell), N. viridis (Newstead); Trionymus newsteadi (Green) *

- Postemesospiracular setae and pores restricted to a group posterior to each mesothoracic spiracle. Lateral finger-like processes on each side of ventral opening of style either present (although maybe small) or absent............. Antonina crawi Cockerell, A. purpurea Signoret; Chaetococcus turanicus Borchsenius; Chorizococcus rostellum (Lobdell); Dysmicoccus vaccinii Miller & Polavarapu; Formicococcus ireneus (De Lotto); Maconellicoccus hirsutus (Green); Paracoccus glaucus (Maskell) ( Fig. 10), Planococcus citri (Risso), P. dioscoreae Williams, P. kenyae (Le Pelley); Pseudotrionymus multiductus (Beardsley); Spilococcus mammillariae (Bouché); Trabutina elastica Marchal ( Fig. 12); Trionymus bocheri Gertsson & Hodgson; T. heterelymus Gertsson & Hodgson, T. thulensis Green; Vryburgia amaryllidis (Bouché) *

Note: *The males of these groups of taxa are morphologically very similar, differing in character-states of unknown taxonomic significance. For species identification within these groups, see mainly Afifi (1968) (and perhaps Beardsley, 1960). §The description and illustration of the macropterous male of Mirococcus ( Polystomophora) ostiaplurimus in Vinis & Kozár (1981) is poor.

Subfamily Phenacoccinae Šulc 1944: 152.

Introduction. This subfamily is considered to be sister to the Pseudococcinae ( Downie & Gullan 2004; Hardy et al. 2008; Kaydan et al. 2015; Danzig & Gavrilov-Zimin 2014). Few adult males have been described in detail but those that have been described are listed in Appendix A. Males in this subfamily are basically similar to those of the Pseudococcinae but differ as in the key above. Some of these character states are similar to those found on adult males of Rhizoecidae, which has been associated with Phenacoccinae by some authors, e.g., Hardy et al. 2008. As examples of phenacoccine mealybugs, illustrations of Phenacoccus solenopsis Tinsley and Phenacoccus sp. (off Pinus halepensis, Evia, Greece, 22.viii.2006, S. Gounari) are here illustrated ( Figs 8, 9).

Subfamily Pseudococcinae Cockerell 1905

Introduction. Quite a few adult males of this subfamily have been described in detail (see Appendix A), starting with Theron 1958, followed Beardsley (1960, 1962, 1964) and Afifi 1968; Afifi & Kozsztarab 1967; Afifi et al. 1976). For more recent studies, see Appendix A. Almost all of these males are macropterous and of rather uniform structure and fall within the diagnosis above. In addition, Williams and Miller (1999) described the adult male of Quadrigallicoccus lauracearum Williams & Miller. This latter species is gall inducing and, although the male is an otherwise typical macropterous male, the abdomen and particularly the penial sheath have become considerably elongated, presumably to assist in fertilization of the female within the gall. This abdominal and/or penial sheath extension is typical of many males of gall-inducing Eriococcidae ( Hodgson & Miller 2010; Semple et al. 2015).

Examples of macropterous adult male pseudococcine mealybugs are illustrated in Figs 10–12, e.g., Paracoccus glaucus (Maskell), Trabutina elastica Marchal and? Spilococcus atriplex (Cockerell); while an apterous pseudococcine mealybug, Oracella acuta (Lobdell) and an allomyrmococcine mealybug, Promyrmococcus dilli Williams are shown in Figs 13, 14.

For differences between Pseudococcidae and Eriococcidae, see key on p. 19.

ERIOCOCCIDAE Cockerell 1899

Type genus: Eriococcus Targioni Tozzetti, 1868, 726.

Type species: Eriococcus buxi Fonscolombe, 1834, 218.

Introduction. The classification of the taxon here referred to as Eriococcidae has been shown to be problematic.All recent phylogenetic analyses, starting with Cox and Williams (1986), have shown that the taxa included in the Eriococcidae form a non-monophyletic group. The name Eriococcidae has been widely accepted since Ferris (1957b) but, even then, the Kermesidae and Dactylopiidae were frequently included in the concept. Later, Hoy (1963) in his catalogue, included the Beesoniidae, Dactylopiidae, Eriococcidae, Kermesidae and Micrococcidae, while earlier Borchsenius (1949) had included the eriococcids in the Pseudococcidae. More recently, various phylogenetic studies have found that the Beesoniidae, Stictococcidae and Dactylopiidae all fall within the overall concept of Eriococcidae but that the Kermesidae do not. Molecular studies ( Cook et al. 2002, Cook & Gullan 2004, Gullan & Cook 2007) found that these four taxa ( Beesoniidae, Eriococcidae, Dactylopiidae and Stictococcidae) all fall within the overall concept of Eriococcidae and were recovered forming four subgroups within an overall clade referred to as the Eriococcidae, namely: (i) Group A: the acanthococcid group comprising the type species of Acanthococcus, A. aceris Signoret, other eriococcids from both the northern and southern hemispheres plus the family Dactylopiidae Costa; (ii) Group B: a Gondwanan group which includes many species from Australia, South America and New Zealand, many of which feed on Myrtaceae; (iii), Group C: a subclade including Beesoniidae, Stictococcidae and some eriococcid taxa, including Eriococcus buxi (Boyer de Fonscolombe), the type species of Eriococcidae (referred to as the BSE group), and (iv) Calycicoccus Brain from South Africa ( Cook & Gullan 2004, Gullan & Cook 2007). Studies based on the morphology of adult macropterous males ( Hodgson 2002, Hodgson & Hardy 2013) also found that the Beesoniidae, Dactylopiidae and Stictococcidae were recovered within the Eriococcid clade, although the division into the above four subclades was less clear. However, the adult males of taxa currently recognised as families, i.e., Beesoniidae, Dactylopiidae and Stictococcidae, appear to be easily separable from the other “eriococcids” based on their morphology and are dealt with under separate headings below. For a key for the separation of these various taxa, see main key on p. 19. Calycicoccus fell within the Gondwanan Group in Hodgson and Hardy (2013) and is discussed under that group heading below.

Group A. ACANTHOCOCCINE GROUP OF ERIOCOCCIDAE

Introduction. The phylogenetic molecular studies mentioned above suggest that this group includes species in the genera Acanthococcus (as per Kozár et al. 2013), Apiomorpha Rübsaamen, Gossyparia Signoret, Eriochiton Maskell and Ovaticoccus Kloet plus the family Dactylopiidae but not Eriococcus Targioni Tozzetti (as per Kozár et al. 2013) (see Kondo et al. 2006). Eriococcus buxi (Fonscolombe), the type species of Eriococcus, was placed within the BSE group (see under this species below). However, neither the Dactylopiidae nor Apiomorpha fell within the acanthococcine group in Hodgson and Hardy (2013), based on adult male morphology alone, and so these taxa are treated separately below.

Group diagnosis (based on adult males of Acanthococcus, Gossyparia, Eriochiton and Ovaticoccus— Acanthococcidae of Kozar et al. 2013). Body not attenuated; with few setae, mainly hs; loculate pores absent (apart from on some Apiococcus sp.); abdomen parallel sided. Head: simple pores often present on head; ocelli close to or touching post-ocular ridge; ventral simple eyes often approximately as far forward as dorsal simple eyes; preocular ridge usually well developed; striations or concentric ridges on ocular sclerite few and restricted to near each simple eye; interocular ridge absent; postoccipital ridge well developed, with both anterior and posterior arms; genal setae present; antennae 10 segmented; capitate setae generally present on several antennal segments in addition to those on apical segment; antennal segments with both hs and fs. Thorax: prosternal ridge present; prescutum usually with prescutal setae; scutal setae present laterally; scutellum with a large foramen and scutellar setae; postmesospiracular setae absent; metasternum usually with few setae; metaprecoxal ridge well developed and extending medially from metapleural ridge; postmetaspiracular setae usually absent; hamulohalteres usually present; alar lobe present; alar setae present; alar sensoria absent; trochanter with 3 oval sensoria on each side placed in a curve or triangle; fs present or absent on legs; tibia with 2+ spurs; tarsi 2 segmented; claw long and narrow, without a denticle; claw digitules capitate. Abdomen: tergites and sternites usually poorly sclerotised or unsclerotised; fs absent; glandular pouches present on segment VIII; pleural setae present on all segments; caudal extensions absent; ostioles absent; abdominal segment IX fused with penial sheath; all setae on penial sheath quite long; segment IX significantly broader than style; anus usually distinct in centre of segment IX; aedeagus usually curved ventrally.

Two previously undescribed acanthocine males are described and illustrated below, Acanthococcus adenostomae (Ehrhorn) and Eriochiton armatus Brittin. For descriptions of adult males of other 14 species, see Appendix A.

For differences between adult males of species in the acanthococcine group and those of Dactylopius, see under the latter genus below.

Key to the known adult male acanthococcine eriococcids

1. Apterous or brachypterous.............................................................................. 2

- Macropterous........................................................................................ 4

2. Apterous. Antennae short, each segment much broader than long, with both fleshy and hair-like setae. Capitate setae absent. Campaniform sensilla on each trochanter in a line. Loculate pores present near each spiracle. With short fleshy setae on each tibia................................................................. Ovaticoccus adoxus (Ferris) * ( Fig. 18)

- Brachypterous. Other characters not in this combination...................................................... 3

3. Capitate setae on antennae absent............................................. Gossyparia salicicola Borchsenius

- Capitate setae on antennae present................................................. Gossyparia spuria (Modeer)

4. Fleshy setae on legs absent............................................................................. 5

- Fleshy setae on legs present............................................................................ 8

5. Antemesopiracular setae absent. Gena with 4–6 genal setae. Abdominal segment VIII with a long pleural seta, about half as long as glandular pouch setae. Small pores present on head.............................. Gossyparia spuria (Modeer)

- Antemesopiracular setae present. Gena with 3 or fewer genal setae. Other characters not in this combination............ 6

6. Hamulohalteres absent although alar lobe present. Small pores on head absent. Penial sheath about 1.5x as long as basal width...................................................................... Ovaticoccus agavium Douglas

- Hamulohalteres present. Small pores on head present. Penial sheath more than twice as long as basal width............. 7

7. Genal setae absent. Loculate pores absent from dorsum of metathorax............... Eriochiton hoheriae Hodgson 1994

- Each gena with 2 setae. Loculate pores present in position of dorsospiracular setae..... Eriochiton armatus Brittin ( Fig. 17)

8. Fleshy setae on legs very short, only about twice as long as broad. Glandular pouch setae each with a small capitate apex................................................................. Acanthococcus adenostomae (Ehrhorn) ( Fig. 15)

- Fleshy setae on legs much longer, several times as long as broad. Glandular pouch setae each with a pointed or flagellate apex................................................................................................... 9

9. Antemesospiracular setae absent. Abdominal segment VIII with one long pleural seta, about half as long as shortest glandular pouch seta. Sclerotised structures associated with aedeagus complex. Penial sheath broad, less than 2x as long as basal width. Cranial apophysis trifurcate.............................. Acanthococcus melnikensis Hodgson & Trencheva ( Fig. 16)

- Antemesospiracular setae present. Other characters not in this combination...................................... 10

10. Penial sheath about 2.5x longer than basal width. Sternite of abdominal segment VIII with a longitudinal sclerotised bar. Head without pores. Without complex sclerotised structures associated with aedeagus.......... Acanthococcus orariensis (Hoy)

- Penial sheath subequal to or less than twice as long as basal width. Sternite of abdominal segment VIII without a longitudinal sclerotised bar. Head with pores. With some complex sclerotised structures associated with aedeagus................................................................................. Uhleria ( Acanthococcus) araucariae (Maskell)

Note.* Although this species ( Fig. 18) is currently placed in Ovaticoccus Kloet, the adult male differs from that of O. agavium in the structure of the campaniform sensilla on the trochanter, which appear to be in a line and oval to elongate (and typical of the Gondwanan group of eriococcids) rather than in a curve or triangle and roundish as in O. agavium and other acanthococcids. In addition, the antennae of adult male O. adoxus are 9 segmented, each segment much wider than long and with few hs and no caps, extremely similar to those of Pseudomontanococcus martini Kozár & Konczné Benedicty ( Fig. 36), whereas those of O. agavium are 10 segmented, each segment longer than wide and each with numerous hs, fs and caps, typical of the acanthococcid group ( Afifi 1968). It would appear that the placement of adoxus in Ovaticoccus warrants further study.

In addition to the two species described below, the adult male of Eriococcus melnikensis Hodgson & Trencheva is also illustrated (and described in Hodgson & Trencheva, 2008) ( Fig. 16).