Corresponding author: Nate B. Hardy (
Academic editor: Michael Wilson
We revise the genus
Here we revise the genus
Freshly collected specimens were slide-mounted in Canada balsam using the method described in
The morphological terms for
Galls of species of
Galls of species of
Galls of species of
This study draws extensively from the collections of P. J. Gullan (98 records), L. G. Cook (25 records), H. M. Brookes (25 records) and N. B. Hardy with PJG (19 records). The late Helen Brookes (formerly at the Waite Agricultural Research Institute, University of Adelaide, South Australia) acquired and curated a large collection of
In the descriptions, we list the slide-mounted specimens that were examined, but most of these collections have associated dry gall material that is too numerous to record and list. Galls and associated slide-mounted insects are stored in the same collections. For many of the species, one to several slide-mounted specimens listed in the ‘Material examined’ are DNA voucher specimens of either L.G. Cook (e.g., LGC00101 for O. thurgoona) or NBH (e.g., NH104 for O. thurgoona) and all are deposited in ANIC (see below). The reliability of early (1890s to 1960s) identifications of eucalypt host plants is uncertain, but more recent records (1970s to present) are considered reliable. The classification of
Depositories are abbreviated as follows: ANIC, Australian National Insect Collection, CSIRO, Canberra, ACT, Australia; ASCU, Agricultural Scientific Collections Unit, Orange Agricultural Institute, New South Wales, Australia; BMNH, the Natural History Museum, London, UK; BPBM, Bernice P. Bishop Museum, Honolulu, HI, USA; NMV, Museum of Victoria, Melbourne, Australia; NZAC, New Zealand Arthropod Collection, Landcare Research, Auckland, New Zealand; QDPI, Queensland Primary Industries and Fisheries, Indooroopilly, Queensland, Australia; SAMA, South Australian Museum, Adelaide; USNM, the United States National Collection of
The
The key by
1. | Abdomen not tapered; anal ring on ventral body surface, usually invaginated, with ≥ 6 ring setae; fore and mid legs and antennae clearly multi-segmented, although often much reduced; cribriform plates or tight clusters of tubular ducts present or absent on dorsum; margins of posterior abdominal segments usually bearing fleshy projections that often bear spines or setae; galls always on leaves | 2 ( |
– | Abdomen tapered; anal ring poorly developed, without pores and with ≤ 6 minute setae, never invaginated, at apex of abdomen or on dorsal surface; fore and mid legs and antennae usually appearing unsegmented and always highly reduced; cribriform plates and tight clusters of tubular ducts absent from dorsum; marginal fleshy projections absent from abdomen (4 spine-tipped fleshy projections on abdominal segment VIII may be present); galls on leaves, stems or occasionally fruits | 7 ( |
2. | Posterior abdominal margin without fleshy projections; fore and mid legs well developed, all leg segments distinct | 3 |
– | Posterior abdominal margin with fleshy projections, often as small segmental lobes; fore and mid legs reduced, some leg segments fused | 4 |
3. | Antenna with 5 fleshy setae; dorsal cribriform plates heavily sclerotised, some on margin composed of > 10 ducts; hind tibia straight; hind tarsus similar to mid and fore tarsi, not expanded | |
– | Antenna with 4 fleshy setae; dorsal cribriform plates weakly sclerotised, none composed of > 10 ducts; hind tibia with distal area of lateral margin concave; hind tarsus expanded, conspicuously larger than mid and fore tarsi | |
4. | Marginal fleshy projections present on abdominal segments anterior to abdominal segment VII, each projection bearing 1–2 blunt or conical spines; dorsal cribriform plates absent | 5 |
– | Marginal fleshy projections restricted to abdominal segment VII and VIII, each projection bearing > 2 spines; dorsal cribriform plates present | 6 |
5. | Cuticle on dorsal head surface rugose and sclerotic; marginal projections on each abdominal segment, with each projection bearing conical spine(s); microducts absent; marginal fringe of elongate setae absent | |
– | Cuticle on dorsal head surface not differentiated from rest of dorsum; marginal projections restricted to abdominal segments IV to VIII or V to VIII, with spines on projections blunt; microducts present; marginal fringe of elongate setae present | |
6. | Dorsal setae much smaller than marginal setae; marginal fringe of close-set conical setae distinct; eye dorsad of marginal fringe | |
– | Dorsal setae about as large as marginal setae; marginal fringe present but not pronounced; eye on margin | |
7. | Dorsum with shield of rugose sclerotic cuticle with 3 deep invaginations along midline; tubular ducts absent | |
– | Dorsal derm membranous, or weakly sclerotic, without shield or deep invaginations; tubular ducts present or absent | 8 |
8. | Dorsum or margin with humps or other evaginations | 9 |
– | Dorsum and margin without humps or evaginations | 13 |
9. | Eyes absent, replaced by a pair of fleshy evaginations | 10 |
– | Eyes present, not mounted on fleshy evaginations | 11 |
10. | One large papilliform evagination present on each side of head on anterior margin, each evagination > length of hind tarsus; other fleshy evaginations absent; coxa conical | |
– | Papilliform evaginations on anterior margin of head < length of hind tarsus; additional evaginations present along dorsal midline plus thoracic submargin; coxa cylindrical | |
11. | Dorsum with three large humps, dorsal setae each mounted on swollen base; eye highly convex, base parallel-sided and perpendicular to body surface; hind tibia curved | |
– | Dorsal humps variable, dorsal setae without swollen bases; eye without parallel-sided base; hind tibia straight | 12 |
12. | Macrotubular ducts present on dorsum; small humps present along midline on thorax and anterior abdominal segments; eye small (20–50 µm wide) | |
– | Macrotubular ducts absent from dorsum; dorsum of mature females dominated by massive humps; eye large (60–85 µm wide) | |
13. | Anal area with 4 stout spines; anal ring sclerotic, sometimes appearing horseshoe-shaped, with 6 fine setae | 14 |
– | Anal area without 4 stout spines; anal ring poorly developed, unsclerotised | 15 |
14. | Fleshy evaginations present immediately caudad of each spiracle; eye large (30–65 mm wide); small spines present along margin of head | |
– | Fleshy evaginations near spiracles absent; eye small (15–25 mm wide); spines absent from head margin | |
15. | Hind tibia and tarsus fused, hind claw absent, tibia and tarsus forming robust sword-shaped segment; tubular ducts absent | |
– | Hind tibia and tarsus separate, hind claw reduced; tubular ducts present | 16 |
16. | Stout spines present in band extending from anterior head margin to posterior spiracles; medial portions of dorsum with numerous conical fleshy projections, each with sclerotic spine at apex, several spines with ante-apical hair-like bristle | |
– | Stout spines absent; dorsum without conical fleshy projections bearing spines | 17 |
17. | Eye large (45–123 mm wide); marginal setae short (longest setae ca 40 mm long), not forming conspicuous fringe | |
– | Eye small (20–45 mm wide); marginal setae long (longest setae ca 150 mm long), forming conspicuous fringe |
Galls on leaves; each typically globular, conical or hemispherical, rarely pit-like; with small circular, slit-like or fissured orifice/opening often on adaxial (upper) surface, but surface often difficult to determine in mature isobilateral leaves and galls typically opening on same surface on any one leaf. Body outline circular to elliptical; dorsum usually smaller than venter, especially at maturity, with whole of dorsum, a sclerotised part of it, or abdominal apex plugging gall orifice; body of mature female usually tightly fitting gall cavity. Abdomen not tapered. Vulva and anal opening ventral on posterior abdomen, with vulva between abdominal segments VII and VIII. Eyes on margin or dorsad of it. Antennae reduced, with ≤ 7 segments, segmentation often indistinct. Pair of broad frontal lobes posteromedial of antennae. Oral lobes membranous to sclerotic, often forming large circular pad around mouthparts. Tentorial box with aliform anterior extensions. Fore and mid legs varying from well developed to highly reduced, with some segmentation always apparent. Hind legs, on posterior of metathorax, sometimes appearing to be on anterior abdomen, large, always well developed, often elongate; ratio of length of trochanter + femur / length of tibia + tarsus approximately = 1:1; tibia-femur articulation functional; translucent pores present on at least tibia; trochanter with 2 or 3 campaniform sensilla on each side; claw digitules and tarsal digitules either well developed or highly reduced to absent. Anal opening surrounded by anal ring with ≥ 6 (range 6–20) setae; anal ring may be invaginated. Posterior abdominal segments usually with marginal fleshy projections (absent in
Antenna 10-segmented. Abdomen not elongated. Gland pouches present, each with pair of setae.
Anterior margin of head incised at midline. Each spiracle with one trilocular pore next to opening. One submedial longitudinal row of dorsal setae on each side of body. Antennae 4-segmented, with 4 fleshy setae on apical segments.
The genus name is a Latinised combination of the Greek words opisthen, meaning behind, and skelos, meaning leg, and clearly refers to the long hind legs of the adult female because
(
Male. On leaf. Similar to gall of female but smaller and more cylindrical, 2–3 mm high, basal diameter 1.5–2.0 mm, with circular opening and truncate apex ca 1 mm across (
(
Dorsum. Dorsal shield much smaller than venter in old specimens, of variably sclerotic cuticle 1.1–2.0 mm long, 1.0–2.7 mm wide, clearly delineated by marginal fringe of close-set spinose setae, each seta 25–70 µm long. A pair of fleshy caudal projections on each side of body; each medial projection with ca 4 spinose setae, each lateral projection (probably marginal lobe of abdominal segment VII) with usually 3 spinose setae. Dorsal setae mostly small and robust, 8–23 mm long, scattered over dorsum, a few large spinose setae similar to those in marginal fringe sometimes occur along submargin. Macrotubular ducts occurring in clusters of 1–25 ducts set within heavily sclerotic cribriform plate; shaft of each duct short, < 5 µm long; number of cribriform plates on dorsum varying from ca 10 to ca 40 on each side of body. Microtubular ducts absent. Quinquelocular pores absent.
Venter. Oral lobes membranous. Setae 8–100 mm long, in transverse row or band across each abdominal segment, metathorax, and mesothorax, along margin of head, pro- and mesothorax; setae dense posterodorsal of anal ring. Macrotubular ducts absent. Quinquelocular pores 5–6 µm in diameter, in clusters around each spiracle and along margin of thorax and anterior abdominal segments.
Adult female of
Holotype of
Paratypes: AUSTRALIA: Victoria: 13 adult females: same data as holotype (11 in ANIC, 1 in BMNH, 1 in USNM).
AUSTRALIA: New South Wales: 4 adult females: ex leaf galls,
The adult female of
Galls of adult females of
This species is named in honour of the late Dr. Jack Beardsley, who was an expert on the systematics of parasitic wasps and scale insects. He spent most of his entomological career at the University of Hawaii, but began studies on Australian gall-inducing eriococcids during a research sabbatical spent in Victoria in 1971–1972. He was interested especially in pit-inducing and cryptic bark-living eriococcids and amassed a substantial collection from Victoria, including some specimens of
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Male. On leaf. Gall opening oblong, surrounded by a slightly raised lip (
(
Dorsum. Head with shield of rugose, sclerotic cuticle, 600–1000 µm long, 450–1100 µm wide; each side of abdomen with segmental pairs of marginal spines, each spine at end of fleshy projection, these larger on posterior segments; each thoracic segment with a few similar structures along the margin; marginal fringe of elongate setae absent. Derm densely beset with microtrichia. Dorsal setae minute, 4–8 mm long; scattered over dorsum. Macrotubular ducts 12–15 µm long, rim of dermal orifice 5 µm wide; ducts in a transverse row across each body segment. Microtubular ducts absent. Quinquelocular pores absent.
Venter. Oral lobes membranous. Ventral setae 5–115 mm long, in transverse row across each abdominal segment, a medial to submedial cluster on each thoracic segment, scattered along margin, longer setae found in medial areas. Macrotubular ducts similar to those on dorsum, scattered along margin, in a transverse row across each of abdominal segments II–IV. Quinquelocular pores 5 mm in diameter, in cluster around each spiracle, along margin of abdomen, in a transverse band across each posterior margin of abdominal segments IV–VIII.
Adult female of
Lectotype of
Paralectotypes: AUSTRALIA: Victoria: 1 adult female, same data and slide as lectotype (ASCU); dry galls (6 of female and 4 of male), all parasitised or empty, with printed label: “3046 E / GALL MAKING COCCIDS, /
The lectotype and associated paralectotype are in poor condition and were mounted from the spirit collection in ASCU by PJG in 1985; additional females, in poor condition, remained in the vial; no original WWF slide-mount was located.
AUSTRALIA: New South Wales: 5 adult females, 5 second-instar females: ex galls on leaves,
Adult females of
(
Male. On leaf. Gall conical (
(
Dorsum. Delineated by fringe of spinose setae along margin, each seta similar to those on rest of dorsum; cauda composed of two truncate fleshy projections on each side of body, each bearing ca 4–6 spinose setae. Derm variously sclerotic, densely beset with minute papillae. Dorsal setae spinose, 20–138 mm long, with stout, or swollen bases. Microtubular ducts absent. Quinquelocular pores absent.
Venter. Oral lobes membranous. Venter hirsute, each seta 23–163 mm long, absent from medial areas of head and pro- thorax, plus antero-medial portion of mesothorax. Macrotubular ducts 10–12 µm long, dermal orifice with rim 5 µm wide; ducts restricted to abdomen, scattered. Quinquelocular pores 5–8 mm in diameter; clustered around each spiracle, and on medial areas of abdomen.
(
Dorsum. Derm membranous. Dorsal setae ca 3 mm long; arranged in submedial longitudinal row on each side of body, 1 seta on each side of head, each thoracic segment, and each or any of abdominal segments I–III and V. Microtubular ducts 4 µm long, on each side of body: 1 duct on submargin of each side of each thoracic segment plus usually each of abdominal segments I, V and VIII. Marginal setae cylindrical, with hemispherical base with diameter > width of cylinder, distal edge of base flat, each marginal seta 2–5 µm long; each side of body with ca 6 setae between midline and eyes, 9 on prothorax, 5 on mesothorax, 3 on metathorax, 2 on each of abdominal segments I–VII, and on abdominal segment VIII 1 lateral and 2 medial of apical seta (these 3 probably homologous to anal lobe setae), with setae medial of apical seta robust, each with truncate apex.
Venter. Setae hair-like, 2–23 mm long, each side of body with 3 setae medial of scape, 1 seta medial of each coxa, 3 longitudinal rows on abdomen, each row with 1 seta on each of abdominal segments II–VII; suranal and ventral lobe setae hair-like, each 18–25 µm long. Trilocular pores 2 µm in diameter, 1 pore near each spiracle.
Adult female of
First-instar nymph of
AUSTRALIA: Australian Capital Territory: 3 adult females:
Adult females of
(
Male. Not known.
(
Dorsum. Delineated by fringe of elongate setae, each seta 50–180 long, with blunt to truncate apex. Derm membranous. Dorsal setae minute, 5–9 mm long, scattered. Macrotubular ducts in two forms: (i) singly or in clusters of 2–8 ducts, cuticle of each cluster variously sclerotic, each duct short, ca 5 µm long, inner ductule not detected, scattered over dorsum, dense on head, posterior abdominal segments and margin of anterior abdominal segments and thorax; and (ii) single, larger ducts, 14–16 mm long, with dermal orifice with a rim 3.5–5.0 mm wide, scattered over dorsum, scarce or absent from posterior abdominal segments. Microtubular ducts absent. Quinquelocular pores absent.
Venter. Oral lobes membranous. Setae 13–103 mm long, in a transverse row across each abdominal segment, scattered along margin of head and thorax, on medial areas of meso- and metathorax, and a transverse band of elongate (up to 170 µm long) setae posterior of frontal lobes. Macrotubular ducts with distal (near vestibule) end of shaft constricted, about same size as larger ducts on dorsum. Quinquelocular pores 5 µm in diameter, overlapping in distribution with ventral setae, with a cluster around each spiracle, but most dense medially on posterior abdominal segments.
Adult female of
Holotype of
Paratypes: AUSTRALIA: New South Wales: 2 adult females: same data as holotype, NH104, LGC00101 (ANIC); 14 adult females, 1 second-instar female, 27 first-instar nymphs: ex galls on leaves,
Adult females of
Live eggs of
The species name is taken from the type locality. It is a noun in apposition.
(
Male. Not known.
(
Dorsum. Delineated by unbroken fringe of moderately dense setae, with approximately 140–170 moderately slender bluntly rounded or minutely capitate setae per side, setae each about 80–165 µm long on abdomen, somewhat shorter anteriorly, 65–100 µm long. Derm membranous. Dorsal body setae sparsely scattered, short (ca 7–15 µm long) and stout, with acute apex. Macrotubular ducts of one kind and size: short, ca 4–7 µm long, with a rim 3–4 µm wide; sparsely scattered singly or in pairs on central part of dorsum, but aggregated into distinct groups on peripheral areas, aggregations increasing in size toward margin, outer ones mostly containing about 10–25 ducts, maximum 30; duct aggregations borne on small, roughly oval to circular, sclerotised tubercles (cribriform plates). Microtubular ducts absent. Quinquelocular pores absent.
Venter. Oral lobes membranous. Ventral body setae sparse, mostly 50–100 µm long, up to 150 µm long on head behind frontal lobes, shorter (20–50 µm long) in lateral pore band. Macrotubular ducts slender, inner portion dilated, 15–17 µm long, 2.0–2.5 µm wide, with poorly sclerotised rim 2.5–3.5 µm in diameter; ducts distributed in a moderately dense peripheral band, ducts distributed evenly across abdominal segments III–VI, but apparently absent on segments immediately in front of and behind vulva where disc pores are most numerous. Quinquelocular pores 5 µm in diameter, mostly confined to abdominal segments V–VIII anterior to anal ring, but a few (12–20) associated with each spiracle.
Adult female of
Holotype of
Paratypes: AUSTRALIA: Victoria: 3 adult females (1 slide), 6 first-instar nymphs (including NH205), 7 embryos, 1 leaf with vacated pit galls: insects from under bark of
Adult females of
The adult female of
The holotype of
There is only minor variation between the adult female holotype from New South Wales and the three paratype females from Victoria; in particular, there are fewer clusters of macrotubular ducts on the dorsal submargin of the Victorian collection.Etymology
The species name is a manuscript name of the late J. W. Beardsley who was describing the species as part of a revision of
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Male. Unknown.
(
Dorsum. Delineated by marginal fringe of elongate (up to 300 µm) flagellate setae, fringe extending from head to abdominal segment IV, margin of each of abdominal segments V–VIII with pair of blunt sclerotic projections at end of short fleshy lamella. Derm densely beset with microtrichia. Dorsal setae flagellate, 15–148 mm long; scattered over dorsum. Macrotubular ducts 14–15 µm long, dermal orifice opening with rim 5 µm wide; ducts in a transverse row across each abdominal segment, scattered over head and thorax. Microtubular ducts 5 µm long, dermal orifice with rim 2 µm wide; distributed evenly over dorsum Quinquelocular pores absent.
Venter. Ventral body surface greatly expanded, posterior abdominal segments only visible from dorsal aspect. Oral lobes membranous. Ventral setae 13–250 mm long, in transverse row across each body segment, scattered along margin. Macrotubular ducts similar to those on dorsum, scattered along margin, in a transverse row across each of abdominal segments II–V. Quinquelocular pores 5 mm in diameter, in a cluster around each spiracle, scattered along margin, in a transverse row across each abdominal segment, most dense around vulva.
Adult female of
Adult female of
Holotype of
Paratypes: AUSTRALIA: South Australia: 6 adult females, 24 first-instar nymphs: same data as holotype (ANIC).
Adult females of
The species name is a combination of the Latin words ungula, meaning hoof, and finis, meaning end or boundary. It refers to the blunt sclerotic projections at the apex of each marginal lamella. The name is a noun in apposition.
This species was described briefly by
Adult female and associated gall. Galls on leaves and stems; each either globular, hemispherical, conical, mammiform or thorn-like, never pit-like; with orifice usually slit-like, sometimes circular to oblong; if galls on leaves, orifice on either abaxial or adaxial surface, depending on
Adult male. Antenna 9-segmented. Abdomen tapered, elongate in some species. Gland pouches and associated setae absent. Detailed description given by
In contrast, the known adult males of
First-instar nymph. Anterior margin of head incised at midline. Each spiracle with one trilocular pore next to opening. One submedial longitudinal row of dorsal setae on each side of body. Antennae 3- or 4-segmented, with 4 fleshy setae on apical segment(s). The known first-instar nymphs of
The genus name is a Latinised combination of the Greek tany- meaning long (from the word tanyo meaning stretch out) and skelos meaning leg, and refers to the long hind legs of the adult female. Our new name is feminine as for
Fuller’s (1897: 1346) original description of
(
Male. Not known.
(
Dorsum. Derm membranous. Dorsal setae robust, often slightly curved, 15–138 mm long; dense over head and thorax, forming transverse band of longer setae on each abdominal segment. Macrotubular ducts 10–15 mm long, dermal orifice with a rim 5 mm wide; in transverse row on each of abdominal segments III–VI. Microtubular ducts absent. Quinquelocular pores 6–8 µm in diameter, scattered over dorsum.
Venter. Derm on abdomen with bands of microtrichia. Setae 15–150 mm long, in a transverse row across each abdominal segment, dense along margin and submargin of head and thorax, medial cluster on mesothorax and metathorax. Macrotubular ducts similar to those on dorsum, in sparse transverse rows across each of abdominal segments III-VI. Microtubular ducts absent. Quinquelocular pores similar to those on dorsum, similar in distribution to ventral setae.
Type material: AUSTRALIA: Western Australia: 3 syntype galls of females, dry and empty (2 dissected open, third gall parasitised), on 2 leaves, all on pin-mount, 2 labels: “
AUSTRALIA: South Australia: 4 first-instar nymphs: ex galls on leaves,
The adult female of
We have examined all available type specimens of Froggatt’s
(
Male (
(
Dorsum. Derm variously sclerotic, beset with small papillae; medial portions of thorax and abdomen with numerous spines each born on a fleshy protuberance, each spine with ante-apical seta, spines diminishing in size caudad. Dorsal setae flagellate, 20–180 mm long; scattered over dorsum. Macrotubular ducts absent. Microtubular ducts absent. Quinquelocular pores absent.
Venter. Derm with microtrichia on abdominal segments. Oral lobes sclerotic, forming large circular feeding pad. Spines similar to those on dorsomedial areas of thorax and abdomen found in submarginal row extending from anterior margin of head to posterior spiracles. Ventral body surface extremely hirsute, setae similar to those on dorsum, flagellate, 30–160 mm long. Macrotubular ducts 12–15 µm long, dermal orifice with rim 5 µm wide; on medial areas of anterior abdominal segments only. Quinquelocular pores large, 7–10 mm in diameter; small cluster around each spiracle.
Adult female of
Lectotype of
Paralectotypes of
Lectotype of
Paralectotype of
In the BMNH, there are also two collections of dry galls labelled as “Opisthoscelis convexa n. sp.” and apparently from WWF (one from Dandenong, Victoria, and one from Sydney, N.S.W.). These appear to have been sent to London in 1922, prior to Froggatt’s description of this species. In 1985, PJG dissected dry galls from the Dandenong collection and slide-mounted two adult females and some first-instar nymphs (listed below). Although WWF presumably identified these as belonging to his new species, with the manuscript name
AUSTRALIA: Australian Capital Territory: 6 adult females: ex galls,
Adult females of
(
Male. Not known.
(
Dorsum. Derm weakly sclerotised, densely beset with small papillae; a conical to papilliform protuberance medially on each thoracic segment and on abdominal segment I, and a similar protuberance on each submargin of thoracic segments I and II, smaller protuberances may be present on each submargin of anterior abdominal segments. Dorsal setae flagellate, 25–212 mm long; arranged in a transverse row across each abdominal segment, scattered over surface of head and thorax; setae increasing in length caudad. Macrotubular ducts 6–12 mm long, dermal orifice with rim 4–5 mm wide; in transverse row across each abdominal segment, scattered over head and thorax. Microtubular ducts absent. Quinquelocular pores absent.
Venter. Derm with microtrichia on abdomen. Oral lobes membranous. Setae 68–200 mm long, in a transverse row across each abdominal segment, along margin of head and thorax. Macrotubular ducts absent. Quinquelocular pores large, 8–10 µm in diameter, similar in distribution to ventral setae.
Adult female of
Holotype (here designated): AUSTRALIA: Queensland: 1 adult female (2.4 mm long, 2.0 mm wide): ex conical gall on leaf,
Paratypes: AUSTRALIA: Queensland: 2 adult females: ex leaf gall,
Adult females of
The species name is derived from the Latin word grallator, meaning ‘stilt walker’, in reference to the extremely long, slender legs of the adult female. The name is a noun in apposition.
We have synonymised
(
Male. On stem and either leaf surface (
(
Dorsum. Derm membranous in young females, older specimens may have clusters of small circular sclerotic regions on margin and submargin of head and thorax, or broad areas of sclerotic cuticle on medial portions of head and thorax. Dorsal setae robust, 10–125 mm long; arranged in a transverse band across each body segment and scattered along margin, longer setae present on abdominal segments. Macrotubular ducts 12 mm long, dermal orifice with rim 5 mm wide; restricted to posterior abdominal segments. Microtubular ducts absent. Quinquelocular pores, 7–11 µm in diameter, scattered across posterior abdominal segments.
Venter. Setae 20–165 mm long, in a transverse row across each posterior abdominal segment plus a cluster extending from each spiracle to margin, a few mesad to each metacoxa, otherwise ventral body surface naked. Macrotubular ducts absent. Quinquelocular pores large, 7–11 mm in diameter, similar in distribution to ventral setae.
Adult female of
Lectotype of
Paralectotypes of
Lectotype of
Paralectotypes of
AUSTRALIA: New South Wales: 1 adult female: ex dry stem gall,
Adult females of
In the original description,
(
Male (
(
Dorsum. Derm membranous in young females, sclerotic in older specimens. Dorsal setae ranging from minute and conical to long and flagellate, 20–175 mm long; arranged in a transverse row or band across each body segment, longer setae present on abdominal segments and forming fringe along entire body margin. Macrotubular ducts 12 mm long, dermal orifice with rim ca 5 mm wide; ducts restricted to posterior abdominal segments. Microtubular ducts absent. Quinquelocular pores large, 7–11 mm in diameter, present on posterior abdominal segments.
Venter. Setae flagellate, 20–165 mm long, in a transverse row across each posterior abdominal segment plus a cluster extending from each spiracle to margin, otherwise ventral body surface naked. Macrotubular ducts absent. Quinquelocular pores similar to those on dorsum, similar in distribution to ventral setae.
Adult female of
Lectotype of
Paralectotypes: AUSTRALIA: New South Wales: 1 adult female: same data as lectotype (ASCU); 1 adult female (slide perhaps from collection of W.M. Maskell since part of label in his handwriting): “B.M. 1945.121 /
AUSTRALIA: New South Wales: 3 adult females: ex galls,
Adult females of
(
Male. On stem; similar to gall of female but without apical cone; gall opening irregularly shaped.
(
Dorsum. Dominated by 3 or 4 large humps. Derm lightly sclerotised, densely beset with minute papillae, these diminishing in size caudad and are replaced by microtrichia on abdomen, cuticle on humps may be more heavily sclerotised than elsewhere. Dorsal setae slender, 7–53 mm long; in transverse row across each abdominal segment, scattered on head and thorax, longest setae on posterior abdominal segments. Macrotubular ducts and microtubular ducts absent. Quinquelocular pores 7–8 µm in diameter, on posterior abdominal segments.
Venter. Oral lobes membranous. Setae as on dorsum, 12–80 mm long, in a transverse row across each abdominal segment, plus a few scattered along margin of head and thorax. Macrotubular ducts absent. Quinquelocular pores similar to those on dorsum, in a transverse row across each abdominal segment plus a few clustered around each spiracle, absent elsewhere.
Adult female of
Holotype (here designated): AUSTRALIA: South Australia: 1 adult female (1.8 mm long, 1.2 mm wide), on slide with 2 other females, holotype closest to data label: ex gall on stem,
Paratypes: AUSTRALIA: South Australia: 2 adult females, on same slide as holotype (ANIC); 6 adult females: ex stem galls,
Adult females of
The original labels on the slides from two of the three collections of
The species name is derived from the Latin word gibber, meaning hump on the back. This species has impressively large dorsal humps. The name is a noun in apposition.
Female. On leaves. Gall a circular raised area, up to 5 mm across and 1–2 mm high; opening slit-like.
Male. Not known.
(
Dorsum. Derm weakly sclerotised, densely beset with small papillae. Dorsal setae flagellate, 50–165 mm long; arranged in a transverse band across each abdominal segment, densely scattered over surface of head and thorax; setae increasing in length caudad. Macrotubular ducts 15 mm long, dermal orifice with rim ca 7 mm in diameter; in transverse row across each abdominal segment, scattered over marginal and submarginal areas of head and thorax. Microtubular ducts absent. Quinquelocular pores absent.
Venter. Oral lobes membranous to sclerotic, forming circular feeding pad around mouthparts. Setae 63–190 mm long, in a transverse band across each abdominal segment plus across meta- and mesothorax, and along margin of head and prothorax. Macrotubular ducts absent. Quinquelocular pores very large, ca 10 µm in diameter, on abdominal segments and medial area of mesothorax.
Adult female of
Holotype (here designated): AUSTRALIA: Queensland: 1 adult female (2.0 mm long, 1.5 mm wide): ex gall on leaf,
Paratypes: AUSTRALIA: Queensland: 5 adult female: same data as holotype (ANIC).
Adult females of
The species name is formed from the Latin word mollis, for soft, and cornuta, for horn. It refers to the pair of fleshy projections on the anterior surface of the head. The name is a noun in apposition.
When
(
Male (
(
Dorsum. Derm membranous. Dorsal setae ranging from minute and conical to long and flagellate, 8–95 mm long; arranged in a transverse row or narrow band across each body segment, longer setae present on abdominal segments and along margin. Spinose seta, 5–10 µm long, found on margin of each posterior abdominal segment, absent from margin of anterior abdominal segments, head and thorax. Macrotubular ducts 15 mm long, dermal orifice with a 5 mm wide rim; in transverse row across each abdominal segment, scattered over thorax, absent from head. Microtubular ducts absent. Quinquelocular pores 5–7 µm in diameter, scattered over dorsum.
Venter. Setae as on dorsum, each 12–175 mm long, in a transverse row or narrow band across each abdominal segment as well as meta- and mesothorax, along margin and submargin of head and prothorax. Macrotubular ducts similar to those on dorsum, restricted to submarginal areas of abdominal segments. Quinquelocular pores similar to those on dorsum, similar in distribution to ventral setae; pores often occurring in pairs.
(
Dorsum. Derm membranous. Dorsal setae fine and minute £1 mm long; arranged in submedial longitudinal row on each side of body, 1 seta on each side of head, prothorax, and each of abdominal segments I–VII. Microtubular ducts 4 µm long, each side of body with 1 duct on submargin of each thoracic segment plus each of abdominal segments I and V; also with 1 duct on submedial area of head, each thoracic segment and abdominal segment VIII. Marginal setae sagittate, 2–5 µm long, each side of body with ca 6 setae between midline and eyes, 4 on prothorax, 3 on mesothorax, 2 on metathorax, 1 on each of abdominal segments I–VII, and on abdominal segment VIII 1 lateral and 2 medial of apical seta, these most likely homologous to anal lobe setae.
Venter. Setae hair-like, each 1–20 mm long, each side of body with 3 setae medial of scape, 1 seta medial of each coxa, 3 longitudinal rows on abdomen, each row with 1 seta on each of abdominal segments II–VII; suranal and ventral lobe setae hair-like, each ca 15 µm long. Trilocular pores 3 µm in diameter, 1 pore near each spiracle.
Adult female of
First-instar nymph of
Lectotype of
Paralectotypes: AUSTRALIA: New South Wales: 2 adult female, ca 400 first-instar nymphs & embryos [10 slides], ex dry galls on leaves and stems: same data as lectotype (ASCU).
AUSTRALIA: Australian Capital Territory: 6 adult females: ex galls,
Adult females of
What we believe to be type material of
(
Male. On leaf (probably adaxial surface). Height 1.8–2.3 mm, width 1.6–4.5 mm, length of basal attachment 1.9–4.6 mm. Gall subconical, apex truncate, opening oblong, 0.7–1.9 mm long, 0.2–0.8 mm wide.
(
Dorsum. Derm weakly sclerotised, densely beset with small papillae. Small medial hump present on each thoracic segment, in some specimens additional humps discernable on anterior abdominal segments. Dorsal setae flagellate, 21–202 mm long; arranged in a transverse row across each abdominal segment, scattered over surface of head and thorax; setae increasing in length caudad. Macrotubular ducts 8–11 mm long, 4–5 mm in diameter; in transverse band across each abdominal segment, scattered across head and thorax. Microtubular ducts absent. Quinquelocular pores absent.
Venter. Oral lobes membranous. Setae 25–165 mm long, in a transverse row across each abdominal segment, along margin of head and thorax. Macrotubular ducts absent. Quinquelocular pores large, 8–11 µm in diameter, similar in distribution to ventral setae except absent on head, and with a cluster near each spiracle.
Adult female of
Lectotype of
Paralectotype: AUSTRALIA: New South Wales: 1 adult female: same data as lectotype.
Lectotype of
Paralectotype: AUSTRALIA: Victoria: 2 adult females, 6 slides with hundreds of first-instar nymphs: same data as lectotype (ASCU).
AUSTRALIA: New South Wales: 1 second-instar female:
Adult females of
(
Male. On leaf, usually opening on adaxial surface. Similar to galls of females; gall surface glaucous if on
(
Dorsum. With medial sclerotic shield composed of crowded nodules separated by deep fissures, with a larger deep invagination at posteromedial margin of each thoracic segment. Derm outside of shield membranous, densely beset with minute nodules. Dorsal setae flagellate, 8–50 mm long, in a transverse row across each abdominal segment, scattered over head and thorax. Macrotubular ducts absent. Microtubular ducts absent. Quinquelocular pores 7–9 µm in diameter, restricted to abdominal segment VIII and lateral areas of VII.
Venter. Mature female with venter expanded relative to dorsum, so ventral body margin appears topologically dorsal. Oral lobes membranous. Setae flagellate, 8–100 mm long, a scant few on abdomen, anterior to mid legs, and along margin. Macrotubular ducts absent. Quinquelocular pores similar to those on dorsum, reasonably numerous in a band radiating from each spiracle to body margin, also a few in transverse band on each of abdominal segments V–VIII.
Adult female of
Holotype (here designated): AUSTRALIA: Victoria: 1 adult female (1.35 mm long, 1.17 mm wide): ex flat, circular gall on leaf,
Paratypes: AUSTRALIA: Victoria: 7 adult females, 4 second-instar female, 10 adult males, 2 pupal males: same data as holotype (ANIC); 5 adult females: ex galls,
AUSTRALIA: Victoria: 9 adult females: ex leaf gall,
Adult females of
Galls of females on
Adult males were observed by PJG emerging from their galls on
The species name is derived from the Latin word poculum, meaning cup and tres or tri- mean three. It refers to the three deep invaginations along the dorsal midline. It is a noun in apposition.
Here we synonymise
(
Male (
(
Dorsum. Derm membranous. Dorsal setae minute, 5–8 mm long; arranged in an irregular transverse row across each abdominal segment, scattered over surface of head and thorax. Stout spinose setae, 5–10 µm long, found on margin of each posterior abdominal segment plus along margin of head and prothorax. Macrotubular ducts 13–15 mm long, dermal orifice with rim 5 mm wide; in transverse row across each abdominal segment, scattered over head and thorax. Microtubular ducts absent. Quinquelocular pores 5–7 µm in diameter, scattered over dorsum.
Venter. Membranous lobe-like protuberance immediately posterior of each spiracle, each protuberance having a few slender setae. Setae flagellate, 30–118 mm long, in a transverse row or narrow band across each abdominal segment, and scattered along margin of head and thorax and mesad to each metacoxa. Macrotubular ducts absent. Quinquelocular pores similar to those on dorsum, in transverse band across each of abdominal segments III–VI, a small cluster near each spiracle, a few scattered among setae along margin of head and thorax, pores often occurring in pairs.
Adult female of
Lectotype of
Paralectotypes: AUSTRALIA: New South Wales: 2 adult females, 1 second-instar female: same data as lectotype (ASCU); 2 adult females, numerous first-instar nymphs 5 adult males: ex leaf galls,
Lectotype of
Paralectotypes: AUSTRALIA: Victoria: 2 adult females, 6 slides with hundreds of first-instar nymphs: same data as lectotype (ASCU).
AUSTRALIA: Autralian Capital Territory: 6 adult females, 1 adult male: ex galls,
Adult females of
There were numerous eggs in the cavity of mature galls of females of
(
Male. Not known.
(
Dorsum. Dominated by 3 large, sclerotic and setose humps, 1 on each thoracic segment, each hump 450–910 µm long, 520–790 µm wide. Derm elsewhere membranous or weakly sclerotised. Dorsal setae robust and conical to flagellate, 10–140 mm long; flagellate setae in a transverse row across each posterior abdominal segment, robust setae mounted on small raised base, scattered over surface of head and thorax, dense on thoracic humps. Macrotubular ducts present or absent, duct 12 mm long, dermal orifice with rim 5 mm wide; if present, only 1 to a few present somewhere on abdominal segments III–VI. Microtubular ducts absent. Quinquelocular pores 7–10 µm in diameter, in a transverse row or narrow band across each posterior abdominal segment.
Venter. Oral lobes membranous. Setae hair-like, each 10–130 mm long, in a transverse row across each abdominal segment; a few setae, each on a raised base, scattered along margin of head and thorax. Macrotubular ducts absent. Quinquelocular pores similar to those on dorsum, in a transverse row to narrow band across each posterior abdominal segment plus a few clustered around each posterior spiracle.
Adult female of
Holotype (here designated): AUSTRALIA: Queensland: 1 adult female (2.2 mm long, 2.0 mm wide): ex gall,
Paratypes: AUSTRALIA: Queensland: 6 adult females, same data as holotype (ANIC); 2 adult females:
Adult females of
The species name is a combination of the Latin words villosus, meaning shaggy, and gibber, for hump. It refers to the setose dorsal protuberances that characterise this species. The name is a noun in apposition.
PJG wishes to acknowledge the late Helen Brookes for her support and assistance with research on gall-inducing scale insects. Specimen loans kindly were arranged by the following people: Peter Gillespie (Agricultural Scientific Collections Unit, NSW Department of Primary Industries, Orange, Australia), Rosa Henderson (Landcare Research, Auckland, New Zealand), Jon Martin (The Natural History Museum, London), Gordon Nishida (formerly of the B. P. Bishop Museum, Hawaii) and Douglass Miller and Debbie Creel (USNM