PSEUDOCOCCIDAE Cockerell 1905

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 : 29-34

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.

Figures

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