DIASPIDIDAE Targioni Tozzetti 1868

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 : 142-147

DIASPIDIDAE Targioni Tozzetti 1868

Diaspis Costa 1828b, 453 . Type species: Diaspis calyptroides Costa by subsequent designation: Cockerell 1902a, 58.

Introduction. The family Diaspididae is the largest family within the Coccomorpha and includes 418 genera and 2595 species ( García Morales et al. 2019). They are all characterised by the production of a scale cover composed of secreted wax filaments which are cemented by the anal fluid and include the two larval exuviae. Until recently, the Diaspididae has been divided into six or seven tribes, including a combination of: Diaspidini , Aspidiotini , Parlatorini, Lepidosaphinini, Leucaspidini , Rugaspidiotini and the subfamily Odonaspidinae . However, this classification, although widely accepted, has been seriously questioned ( Takagi 1969, 1997; Morse & Normark 2006; Anderson et al. 2010) and most of the above tribes have been shown to be non-monophyletic. A new classification, based mainly on nuclear, mitochondrial and endosymbiont gene sequences ( Normark et al. 2019) divides the family into 4 subfamilies, namely Ancepaspidinae, Furcaspidinae, Diaspidinae and Aspidiotinae ; divides the Diaspidinae into 2 tribes, the Lepidosaphenini and Diaspidini , and the latter tribe into 3 subtribes, the Diaspidina, Fioriniina and Chionaspidina. The Aspidiotinae are divided into 7 tribes, namely Gymnaspidini, Leucaspidini , Aonidiini, Smilacinini, Odonaspidini , Parlatoriini and Aspidiotini . This classification is followed below.

Most diaspidid males are macropterous but a few apterous species are known. In their molecular study, Gullan and Cook (2007) found the Diaspididae sister to a clade which included the Gondwanan eriococcids and the BSE eriococcids and Yokogawa and Yahara (2009) also found the diaspidids sister to the eriococcids (neither work included the Conchaspididae ). Hodgson and Hardy (2013), based on male morphology, found diaspidids in a clade sister to the Conchaspididae and both families were sister to most other neococcoids. Vea and Grimaldi (2016) found the diaspidids in a clade that included all of the “eriococcids” plus the Dactylopiidae , Pityococcus , Conchaspididae and the Phoenicococcidae .

Quite a lot of male diaspidids have been described in the past: Theron (1958) described male Pseudaulacaspis (Aulacaspis) pentagona (Targioni Tozzetti) and Diaspidiotus bavaricus (Lindinger) ; Ghauri (1962) described the males of 24 species; Nada and Mohammad (1984) described the male of Leucaspis riccae (Targioni Tozzetti) whilst Lambdin et al. (2007) described that of Fiorinia externa Ferris ( Fig. 59 ). Thus, one or more adult males of most of the currently recognised tribes have been described and illustrated. Below is a description of two more Leucaspidini , namely Leucaspis gigas (Maskell) and Labidaspis myersi (Green) , the latter usually assigned to the Parlatoriini but found to be in the Leucaspidini by Anderson et al. (2010).

Family diagnosis based on adult male morphology ( Figs 59–61 ). Body. Abdomen rather parallel-sided, with an extremely long, needle-like penial style; body setae sparse, all short and hs; loculate pores absent. Head: approximately oval, longer than wide, with postocular ridge causing a shallow constriction in side of head; width across genae much wider than rest of head; neck entirely absent; postoccipital ridge very well developed; dorsal head setae few, all hs; mid-cranial ridge present or absent dorsally, generally short ventrally; ocelli present or absent; with two pairs of simple eyes only; ventral eyes approximately below dorsal eyes; genae very broad, not sclerotised or reticulated; genal setae present or absent; ventral head setae few, all hs and restricted to just anterior and mesad to ventral eyes (interocular setae); preocular ridge absent or ill-defined ventrally, short dorsally; ocular sclerite sclerotised but usually not reticulated; antennae quite long, 10-segmented; antennae with a few short hs and some long fs, each much longer than width of antennal segments; apical antennal segment with a capitate seta arising on apex and sometimes 1 or 2 more caps laterally; scape with or without setae. Thorax. Prothoracic setae present or absent; prosternum with a well-developed median ridge; transverse ridge distinct; prescutum convex, with a well-developed prescutal ridge; prescutum without prescutal setae; scutum without a median membranous area; scutum without setae or with a pair just anterior to scutellum; scutellum tubular, usually with a distinct small foramen; scutellar setae present or absent; area delineated by scutellum anteriorly and mesopostnotum posteriorly unsclerotised; postmesospiracular setae absent; basisternum without a strong median ridge or with just a hint of a ridge; furca broad basally, well-developed, with arms at an oblique angle; basisternal setae absent; dorsal part of mesopleural ridge present; suspensorial sclerite present or absent; metepisternum sometimes slightly sclerotised; postmetaspiracular setae absent; metasternal setae generally absent; alar setae absent; alar sensoria absent; alar lobe present; hamulohalteres present; legs fairly setose, mostly hs but tarsi with spur-like setae, sometimes bifurcated; trochanter very long, about 1/2–1/3 length of femur; trochanter sensilla in a line; tibia about 1.5x length of tarsus; tarsal campaniform pore present; tarsal digitules finely capitate; claw digitules capitate or setose; claw without a denticle. Abdomen. Tergites and sternites sometimes sclerotised; with 1 or 2 pleural setae on each side of segments II–VII; caudal extensions sometimes present on segment VII; segment VIII sclerotised and ring-like, but not forming part of penial sheath; glandular pouches absent; segment IX sclerotised and fused to penial sheath; segment IX with 1 pair of long setae; style broad anteriorly, rapidly narrowing into a long, needle-like shaft; with 2 pairs of setae anteriorly; aedeagus long (sometimes longer than penial sheath); basal rod short or perhaps absent.

No males of the following taxa have yet been described: families Ancepaspidinae and Furcaspidinae, or of the following tribes or subfamilies within the Aspidiotinae : Gymnaspidini, Aonidiini, Smilacicolini and Odonaspidina. A “tentative” key for the separation of most of the groups and males of 25 species of Diaspididae can be found in Ghauri (1962). [Note than the Latin names of many of the species studied by him have changed since his study]. The following simplified key follows Ghauri with the addition of a few more recently described species.

Key to adult male Diaspididae (modified after Ghauri, 1962)

1. Ocelli generally absent*. Pronotal ridge absent. Mid-cranial ridge well developed dorsally. Postocular ridge obvious dorsally … Diaspidinae....................................................................................... 2

- Ocelli present laterally just anterior to postocular ridge. Pronotal ridge present. Mid-cranial ridge very short or absent dorsally (except Leucaspidini ). Postocular ridge often not visible dorsally on head … Aspidiotinae ........................... 8

2. Each scape with a long seta ventrally. Each anterior trochanter with an apical spine. Scutum and prescutum without reticulations. Antennae with capitate setae in addition to apical seta. Genal spot* absent. With 2 pairs of genal setae............. 3

- Each scape without any setae. Each anterior trochanter without an apical spine but with lateral teeth (except Prodiaspis ). Scutum and prescutum with distinct reticulations (except Prodiaspis ). Antennae without capitate setae in addition to apical seta. Genal spot* present or absent. With 1 pair of genal setae (except Prodiaspis )...................................... 5

3. Postoccipiral ridge with a median extension posteriorly. With 2 pairs of tentorial pits. Suspensorial sclerites absent … Lepidosaphidini................................................ Lepidosaphes ( L. ulmi (L.); L. pinnaeformis (Bouché))

- Postoccipiral ridge without a median extension posteriorly. With 1 pair of tentorial pits. Suspensorial sclerites present...... Diaspidina.......................................................................................... 4

4. Basisternum with a weak median ridge. Third antennal segment with some hair-like setae in addition to long fleshy setae. Claw digitules dissimilar, with digitules on posterior claws much shorter than those on anterior claws.................................................................................................... Diaspis ( D. boisduvalii Sign. )

- Basisternum without a median ridge. Third antennal segment without hair-like setae in addition to long fleshy setae. Claw digitules similar on all legs.......................... Carulaspis ( C. juniperi (Bouché) ; C. minima (Targioni Tozzett)) .

5. Trochanter of forelegs without teeth laterally. Scutum and prescutum without reticulations. Mid-cranial ridge dorsally without mid-cranial ridge setae. With 1 pair of interocular setae. Antennal setae few and very short, less than width of each segment........................................................................ Prodiaspis tamaricicola (Malenotti)

- Trochanter of forelegs with teeth laterally. Scutum and prescutum with distinct reticulations. Mid-cranial ridge dorsally with mid-cranial ridge setae. With 3 or 4 pairs of interocular setae. Antennal setae abundant and long, much longer than width of each segment........................................................................................ 6

6. Genal spot* absent......................................................................... Fioriniina … 7 - Genal spot* present.. Chionaspidina ( Aulacaspis rosae (Bouché) , Chionaspis salicis (L.), Pinnaspis aspidistrae (Signoret)) ǂ

7. Median ridge of basisternum present. With only 1 pair of genal setae....... Pseudaulacaspis pentagona (Targioni Tozzetti) - Median ridge of basisternum absent. With 2 pairs of genal setae....................... Fiorinia externa Ferris ( Fig. 59 )

8. Mid-cranial ridge divided on dorsal surface of head, each arm extending latero-posteriorly to near each dorsal simple eye, form- ing an inverted V. Postocular ridge on dorsal surface of head distinct … Leucaspidini ............................... 9

- Mid-cranial ridge absent or very short on dorsal surface of head. Postocular ridge on dorsal surface of head rarely visible. 10

9. Antennae quite long, most segments elongate, about 5–6x longer than width...................................................................................... Leucaspis gigas (Maskell) ( Fig. 60 ), L. riccae (Targioni Tozzetti)

- Antennae shorter, rather barrel-shaped, most segments about 3–4x longer than width... Labidaspis myersi (Green) ( Fig. 61 )

10. Scape with a minute hair-like seta on dorsal surface. Apical antennal segment with 1 capitate seta in addition to apical capitate seta. Prosternum with both transverse and median ridges developed. Prescutal and scutellar setae present........................................ Parlatoriini ( P. blanchardi (Targioni Tozzetti) , P. crypta McKenzie , P.?morrisoni McKenzie))ǂ

- Scape without a minute hair-like seta on dorsal surface. Apical antennal segment with 2 capitate setae in addition to apical capitate seta. Prosternum with only median ridge developed. Prescutal and scutellar setae absent … Aspidiotini ......... 11

11. Posterior arms of postoccipital ridge extending laterally almost in a straight transverse line. Anterior arms of postoccipital ridge poorly sclerotised, represented by a sclerite. Cranial apophysis short, not reaching ventral eyes. Dorsal mid-cranial ridge absent................................................................... Diaspidiotus bavaricus (Lindinger)

- Posterior arms of postoccipital ridge extending posterior-laterally, not nearly forming a straight transverse line.Anterior arms of postoccipital ridge strongly sclerotised. Cranial apophysis long, clearly reaching ventral eyes. Dorsal mid-cranial ridge present or absent........................................................................................... 12

12. Dorsal and ventral simple eyes particularly large, with each cornea 17–50 µm wide............................... 13

- Dorsal and ventral simple eyes small, with each cornea 12–21 µm wide......................................... 15

14. Membranous area bordered anteriorly by scutellum and laterally by mesopostnotum, without longitudinal microridges............................................................. Aonidiella orientalis Newstead ; A. aurantii (Maskell) ǂ

- Membranous area bordered anteriorly by scutellum and laterally by mesopostnotum with longitudinal microridges................................................... Aspidiotus nerii (Bouché) ; A. destructor Signoret ; A. excisus (Green) ǂ

15. Flagellar segments of antennae with segments barrel-shaped, each about 2x longer than wide..........................

................................................... Diaspidiotus ostraeiformis (Curtis) , D. zonatus (Frauenfeld) - Flagellar segments of antennae longer, each about 3x longer than width.............. Hemiberlesia cyanophylli (Signoret)

* Ghauri (1962, p. 16), following the views of previous workers, considered that the small dark patch on each gena of some Chionaspidini were “rudimentary” or “larval” eyes. However, as they are on the gena, they are posterior to the postocular ridge whereas, on all other Coccomorpha , the ocelli (larval eyes) are present on the ocular sclerite just anterior to the postocular ridge. It is considered here, therefore, that these spots or patches on the gena on mounted specimens are not “larval eyes”, although their homologies are otherwise uncertain. They are here referred to as “genal spots”. Note that what are referred to as “genal tubercles” occur on some Lecanodiaspididae but whether these are homologous with genal spots is unknown.

ǂ These groups of taxa are all very similar and the significance of the small morphological differences is uncertain.

Figures

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

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

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