(Fig. 1a, b, c). Female. On leaf. Gall opening circular, 0.3–0.4 mm wide, blocked by white wax and/or part of sclerotised posterior dorsum of female. Gall variable, usually 3–5 mm in diameter, side with opening subconical (Fig. 1a) or hemispherically rounded (Fig. 1b), with truncate apex, typically on adaxial surface; leaf glands enlarged.

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 (Fig. 1c), usually on adaxial leaf surface.

(Fig. 3) (n = 47). Body outline circular, length 1.4–3.3 mm, greatest width 1.2–2.8 mm, abdomen not tapered, anal opening ventral, vulva as far cephalad as distal end of hind coxa. Eyes set well away from margin, each 20–60 µm wide. Antennal segmentation indistinct; each antenna 63–120 mm long. Frontal lobes each 110–330 µm long, 83–350 µm wide. Tentorial box 153–590 mm long; labium 73–125 mm long, 65–128 mm wide; pump chamber 17–22 µm long, 20–23 µm wide. Spiracles 63–170 mm long, 35–90 mm wide across atrium. Fore and mid legs small stumps, some segmentation apparent, 68–190 µm long; hind leg with coxa 200–440 µm long, trochanter + femur 260–510 µm long, tibia slightly curved, outer margin concave, 250–410 µm long, tarsus 70–180 µm long; claw and digitules present but reduced; translucent pores dense on both surfaces of hind tarsus, tibia and distal portion of femur; tibia-femur articulation functional. Anal opening 23–55 µm wide; anal ring 48–110 µm wide, bearing 10–16 setae, most anal ring setae with 1 or 2 pores near base; anal ring set within membranous invagination; area between anal ring and dorsal shield sclerotised in some specimens.

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.

Holotype of Opisthoscelis beardsleyi (here designated): AUSTRALIA: Victoria: 1 adult female (2.3 mm long, 2.2 mm wide): ex gall, Eucalyptus goniocalyx, Tallangatta, Mt Granya, 26 May, 1976, PJG (ANIC).

Paratypes: AUSTRALIA: Victoria: 13 adult females: same data as holotype (11 in ANIC, 1 in BMNH, 1 in USNM).

The adult female of Opisthoscelis beardsleyi is most similar to that of Opisthoscelis subrotunda in that both species have dorsal cribriform plates, a marginal fringe of spinose setae, and ≥ 2 spinose setae mounted on each caudal fleshy projection or lobe (these projections poorly developed in some populations of Opisthoscelis beardsleyi). Adult females of Opisthoscelis subrotunda can be recognised by having the dorsal derm densely beset with minute papillae (found in some species of Tanyscelis but no other Opisthoscelis species) and large, spinose dorsal setae (dorsal setae minute in Tanyscelis, or bristle-like in other species of Opisthoscelis).

Galls of adult females of Opisthoscelis beardsleyi may be conical (Fig. 1a) or rounded with either a truncate (Fig. 1b) or convex apex. Adult females vary in the number of cribriform plates and enlarged setae on the dorsal submargin as well as the extent to which the caudal projections are developed. It is possible that the material we have listed under the name Opisthoscelis beardsleyi constitutes multiple species, but we have not been able to detect any clear patterns in morphological variation over geography or host use. Therefore, we have restricted the type material to a single collection. Specimens of Opisthoscelis beardsleyi have been collected only from southeastern Australia, from eucalypt species in the Symphyomyrtus section Maidenaria.

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 Opisthoscelis and Tanyscelis.

(Fig. 1d, e, f).Female. On leaf. Height 2.4–7.0 mm, width 2.5–6.6 mm, length of basal attachment 2.4–4.0 mm. Gall opening slit-like ca 2 mm wide, on abaxial leaf surface. Gall spherical, sometimes with a tapered point (Fig. 1d), opening on opposite side of leaf. Mature female filling gall cavity (Fig. 1e).

Male. On leaf. Gall opening oblong, surrounded by a slightly raised lip (Fig. 1f), on abaxial leaf surface; opposite side of leaf with small globose swelling.

(Fig. 5) (n = 21). Body outline ovate, length 2.3–5.0 mm, greatest width 1.8–4.3 mm; abdomen about as long as head + thorax. Eyespots set well away from margin, each 28–90 mm wide. Antennae 6-segmented, each 290–490 mm long. Frontal lobes may be difficult to discern, each 225–470 µm long, 150–490 µm wide. Tentorial box 285–480 mm long. Labium 75–155 mm long, 100–168 mm wide. Pump chamber 25–28 µm long, 23–30 µm wide. Spiracles 68–195 mm long, 48–85 mm wide across atrium. Fore and mid legs reduced but segmented, 43–160 µm long. Hind legs slender and elongate; coxa 300–590 µm long; trochanter + femur 480–940 µm long; tibia straight, 460–1000 µm long; tarsus 180–280 µm long; claw and digitules present but reduced; few translucent pores scattered on femur, tibia and tarsus; femur-tibia articulation functional. Anal opening ventral, set in shallow membranous invagination, 30–55 µm wide; anal ring plate-like, 63–128 µm wide, with 6–18 setae.

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.

Lectotype of Opisthoscelis serrata (here designated): AUSTRALIA: Victoria: 1 adult female (damaged, ca 3.0 mm mm long, 2.7 mm wide), on slide with another adult female, lectotype farthest from data label; original label: “Opisthescelis[sic] serrata / ♀ galls on Euc. Adult in / spirit. Slide in coll. / Bendigo W.W.F. / Damage in Box 284” ASCTHE101344 (ASCU).

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, / Opisthoscelis serrata, Frogtt. / Female galls upon Eucalyptus sp. / Bendigo, Victoria.” ASCT00004861 (ASCU).

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.

Adult females of Opisthoscelis serrata can be recognised easily by (1) the shield of rugose sclerotic cuticle on the dorsal surface of the head, and (2) the pair of marginal spine-tipped fleshy projections on each side of each abdominal segment. Both of these features are unique amongst Opisthoscelis and Tanyscelis species. Opisthoscelis serrata is known only from eucalypt species in the section Adnataria. There is some variation in the shape and size of the gall of the adult female, which is basically globular. Galls on leaves of Eucalyptus largiflorens and Eucalyptus polyanthemos protrude as a sphere on one side of the leaf, although some galls from leaves of Eucalyptus polyanthemos at Warrandyte have a pointed apex (Fig. 1d). In contrast, galls from the leaves of Eucalyptus melliodora and Eucalyptus microcarpa from near Nagambie protrude equally from both leaf surfaces and the females are smaller than those from most other localities. Females from all galls look similar, although differing in size.

Schrader (1863) described and illustrated Opisthoscelis subrotunda based on insects and galls from an unidentified Eucalyptus sp. collected in New South Wales. Schrader’s collection was destroyed on July 30, 1943, during WWII according to Ew. Weidner in personal communication to PJG (Gullan 1984a). Rübsaamen (1894) described a number of Australian gall-inducing insects, including Opisthoscelis globosa from Eucalyptus in New South Wales, while his address was “Königlichen Museum für Naturkunde zu Berlin”. We contacted Museum für Naturkunde Leibniz-Institut für Evolutions- und Biodiversitätsforschung an der Humboldt-Universität zu Berlin, but were told that there was no Opisthoscelis material from Rübsaamen in that collection, and thus we believe that Rübsaamen’s material also was destroyed during WWII, as suggested by Gullan (1984a). The synonymy of Opisthoscelis subrotunda and Rübsaamen’s Opisthoscelis globosa was proposed by Froggatt (1894a) but not accepted by Hoy (1963); subsequent authors have followed Hoy, although Miller and Gimpel (2000) stated that the status of these two species needed further study. Here we accept Froggatt’s (1894a) synonymy of these two species based on the perfect match of Schrader’s and Rübsaamen’s descriptions and illustrations of the insects and their galls. There is only one species in New South Wales that exactly matches Schrader’s and Rübsaamen’s descriptions. Thus we are certain of the identity of Opisthoscelis subrotunda and consequently have not designated a neotype.

(Fig. 1g, h, i). Female. On leaf. Gall globose (Fig. 1g), surface punctate, leaf glands enlarged on gall. Gall height 4.0–8.7 mm, width 4.2–9.5 mm, length of basal attachment 3.0–6.8 mm; wall of gall at least 2 mm thick in most mature galls. Gall opening variable, slit-like, cruciform (Fig. 1h) or round, sometimes with “lips” projecting, 0.1–2.5 mm wide; opening on abaxial or adaxial surface, but usually on same surface on any one leaf or plant. Mature female fills gall cavity with her abdominal apex directed towards and plugging gall orifice.

Male. On leaf. Gall conical (Fig. 1i), distal margin entire, opening oblong to round, 0.5–2.2 mm wide. Gall height 1.4–4.1 mm, width 1.1–3.6 mm, length of basal attachment 1.3–3.7 mm.

(Fig. 6) (n = 30). Body outline ovate, length 2.1–4.8 mm, greatest width 1.5–4.0 mm; abdomen about as long as head + thorax. Eyespots on margin, each 40–60 mm wide. Antennal segmentation poorly developed; each antenna 50–160 mm long. Frontal lobes sometimes difficult to discern, each 140–280 µm long, 100–240 µm wide. Tentorial box 260–560 mm long. Labium 70–140 mm long, 70–160 mm wide. Pump chamber 26–33 µm long, 28–33 µm wide. Spiracles 90–180 mm long, 50–120 mm wide across atrium. Fore and mid legs reduced but segmented, 115–360 µm long. Hind legs slender and elongate; coxa 370–640 µm long; trochanter + femur 520–950 µm long; tibia 420–1020 µm long; tarsus 110–170 µm long; claw and digitules present but reduced; translucent pores on both surfaces of tibia and tarsus; femur-tibia articulation functional. Anal opening ventral, set in shallow membranous invagination, 38–83 µm wide; anal ring 65–120 µm wide, with 8–16 setae.

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.

(Fig. 7) (n = 4). Body outline elliptical, anterior margin incised at midline; length 288–308 µm, greatest width 180–198 µm. Eyespots on margin, each 10–11 mm wide. Antennae 4-segmented, each ca 75 mm long. Tentorial box 55–63 mm long. Labium 25–28 mm long, 28–33 mm wide. Spiracles 13–19 mm long, 8 mm wide across atrium. Each leg: coxa 23–28 µm long, with 5 setae; trochanter + femur 60–66 µm long, trochanter with 4 setae, femur with 4 setae; tibia 38 µm long, with 4 setae; tarsus 38–40 µm long, with 4 setae; claw 15–18 µm long; tarsal digitules capitate, unequal length, short digitule 23–24 µm long, long digitule 28–33 µm long, claw digitules capitate, each 15–18 µm long. Anal ring 15–16 µm wide, with 6 setae, each ca 13–18 µm long. Apical seta 135–183 µm long.

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.

AUSTRALIA: Australian Capital Territory: 3 adult females: Eucalyptus sp., Canberra, Bruce, 23 Sep., 1972, H. J. Banks (ANIC). New South Wales: 1 adult female: ex leaf gall, Eucalyptus blakelyi, 26 km N of Canowindra, 14 km S of Cudal, -33.41°; 148.69°, 13 Aug., 2004, NBH and PJG, NH21 (ANIC); 13 adult females: ex galls on leaves, Eucalyptus longifolia, 8.5 km SSE Moruya, Bingie Road, -35.97°; 150.12°, 7 Jan., 1992, PJG (ANIC); 5 adult females, 3 second-instar females, 2 first-instar nymphs: ex galls on leaves, Corymbia gummifera, Bundeena, roadside, 14 Feb., 2004, C. A. M. Reid and PJG, NH84 (ANIC); 5 adult females: ex leaf, Eucalyptus longifolia, Congo, 4 Jan., 1992, PJG (ANIC); 2 adult females: ex galls on leaves, Eucalyptus sp., Congo, Congo Road, -35.95°; 150.15°, 8 Jan., 1992, PJG (ANIC); 10 adult females: Sydney (ASCU); 1 adult female: ex leaf gall, Eucalyptus sp., Toolom, near Toolom Falls on edge of Yabbra State Forest, -28.52°; 152.53°, 22 Aug., 2004, NBH and PJG, NH23 (ANIC); 7 adult females (2 slides): Eucalyptus robusta, Sydney Harbour, W. W. Froggatt #1868 (ANIC); 4 adult females (2 slides): Eucalyptus resinifera, Taree, WWF #1941 (ANIC); 1 adult female: Eucalyptus sp., Hay, 1919, WWF #299 (BMNH); 1 adult female: Eucalyptus robusta, Sydney Harbour, 20 Dec., 1929, WWF (BMNH). Northern Territory: 5 adult females, 13 adult males: Eucalyptus camaldulensis var. obtusa, ca 120 km W of Alice Springs, Ormiston Gorge, 29 May, 1977, PJG (ANIC); 4 adult females: ex galls on leaves, Eucalyptus camaldulensis, Serpentine Gorge Nature Park, 27 May, 1992, PJG and P. S. Cranston (ANIC). Queensland: 1 adult female: ex gall, Eucalyptus resinifera, on road from Paluma to Hidden Valley, -19.02°; 146.12°, 30 Oct., 2003, LGC and M. D. Crisp, LGC00066 (ANIC); 4 adult females: Eucalyptus rostrata (now Eucalyptus camaldulensis), Barakula, 13 Apr., 1939 (QDPI); 1 adult female: Eucalyptus rostrata (now Eucalyptus camaldulensis), Barakula, Nov. 1939, No, 784 (QDPI); 5 adult females: Eucalyptus crebra, Clermont, Nov. 1939 (QDPI); 5 adult females: Eucalyptus melliodora or Eucalyptus rostrata (now Eucalyptus camaldulensis), Emu Vale, 8 Feb., 1939, No. SC394 (QDPI); 5 adult females: on leaves, Eucalyptus rostrata (now Eucalyptus camaldulensis), Emu Vale, 15 Mar., 1939 (QDPI); 3 adult females: Eucalyptus propinqua, Monto, 9 Nov. 1938, No. SC375 (QDPI); 3 adult females: Eucalyptus saligna, Pechey, 5 Mar., 1937, INSECOLL 0-067151, 0-067152 (QDPI); 2 adult females and ca 25 first-instar nymphs: Eucalyptus pilularis, Pechey, May 1938, No. SC326 (QDPI); 3 adult females: Eucalyptus tereticornis, Redland Bay, 9 Apr., 1939, No. 785 (QDPI); 1 adult female: Eucalyptus grandis, Tamborine, 20 Jan., 1954, INSECOLL 0-067167 (QDPI). Victoria: 1 adult female: ex gall on leaf, Eucalyptus camaldulensis, 27 km E of Shepparton, nr Midland Hwy, 8 Feb., 2004, PJG, LGC00099 (ANIC); 4 adult females: ex galls on leaves, Eucalyptus camaldulensis, Bundoora, La Trobe University, Wildlife Reserves off Ring Road, -37.72°; 145.05°, 14 Feb., 2005, PJG and NBH, NH43 (ANIC, NMV); 7 adult females: Eucalyptus mcintyrensis (= ? Eucalyptus camaldulensis or a hybrid), Glenthomson, 14 Feb., 1964, HMB, Specimen Index No. 08/64 (ANIC); 1 adult female: ex leaf gall, Eucalyptus camaldulensis, Mildura, River Road near Lock 11, Murray River, -34.17°; 142.16°, 4 Feb, . 2005, NBH and PJG, NH75 (ANIC); 1 adult female: ex globose gall on leaf, Eucalyptus ?camaldulensis, Shepparton, near Broken River, off Lincoln Drive, close to Goulburn Valley Hwy, -36.40°; 145.39°, 30 Jan., 2005, PJG, NH72 (ANIC); 1 adult female: ex globose leaf gall, Eucalyptus ?microcarpa, Shepparton, near Goulburn River, off Tom Collins Drive, -36.39°; 145.39°, 29 Jan., 2005, PJG, NH71 (ANIC). Western Australia: 8 adult: ex galls on leaves, Eucalyptus camaldulensis, Winjana Gorge Nat. Park, bank of Lennard River, -17.42°; 124.95°, 29 Apr., 1992, PJG (ANIC). PAPUA NEW GUINEA: 8 adult females, 20 first-instar nymphs: ex galls on leaves, Eucalyptus alba, Central Province, Sogeri, -9.43°; 147.58°, 10 Apr., 1980, H. Roberts (ANIC, BMNH).

Adult females of Opisthoscelis subrotunda most closely resemble those of Opisthoscelis beardsleyi [see comments under Opisthoscelis beardsleyi]. Specimens of Opisthoscelis subrotunda have been collected from six Australian states or territories as well as Papua New Guinea. This is the only species of Opisthoscelis or Tanyscelis to have been collected outside of Australia. Opisthoscelis subrotunda also appears to have a much broader host range than is typical for species of Opisthoscelis and Tanyscelis. Specimens have been collected from Corymbia in addition to hosts in the Eucalyptus sections Adnataria, Exsertaria, Latoangulatae, and Similares (all in subgenus Symphyomytrus). A common host of this species is the river red gum, Eucalyptus camaldulensis, which has the widest natural distribution of any eucalypt species and seven subspecies (Chippendale 1988; McDonald et al. 2009). The length of the dorsal setae, and the size of the cribriform plates of the adult females of Opisthoscelis subrotunda vary, but in general, the morphology is remarkably homogeneous across samples. Molecular data from collections of fresh material from across the geographic range of the taxon would be required to determine if cryptic species are present.

(Fig. 1j). Female. On leaf. Immature gall (housing yellow nymph with a red dorsal “keel”) shallowly conical to hemispherical on orifice side (Fig. 1j), with gall surface whitish green to reddish; almost hemispherical on opposite side of leaf, tissue reddish green with distinct oil glands on surface. Gall height on orifice side ca 1 mm, ca 3 mm on opposite side, width 4–6 mm. Gall opening on either abaxial or adaxial leaf surface, slit-like, 0.5–1.0 mm long. Mature gall with tissue surrounding orifice brown and necrotic.

Male. Not known.

(Fig. 8) (n = 6). Body ovate, length 2.1–3.4 mm, greatest width 1.5–2.8 mm; margin without lobes or indentations. Eyes on margin, each 38–55 µm wide. Antennae well developed, 7-segmented; each antenna 530–890 mm long; segment I with ca 3 hair-like setae; II with 5–9 hair-like setae; III with 4–8 hair-like setae; IV 4–6 hair-like setae; V with 1–4 hair-like setae; VI with 3 or 4 hair-like setae + 1 fleshy seta; VII with ca 6 hair-like setae + 3 fleshy setae. Frontal lobes each 300–310 µm long, 125–220 µm wide. Tentorial box 200–300 mm long; labium 80–150 mm long, 95–170 mm wide; pump chamber 15 µm long, 15–16 µm wide. Spiracles 100–130 mm long, 50–73 mm wide across atrium. All legs well developed, hind legs larger than fore or mid legs: fore leg with coxa 230–360 µm long, trochanter + femur 345–530 µm long, tibia 235–400 µm long, tarsus 100–140 µm long; mid leg with coxa 310–430 µm long, trochanter + femur 370–580 µm long, tibia 260–450 µm long, tarsus 110–150 µm long; hind leg with coxa 350–500 µm long, trochanter + femur 460–670 µm long, tibia slightly curved, outer margin concave, 380–580 µm long, tarsus 150–240 µm long; each leg with claw 23–40 µm long; translucent pores dense on hind tarsus and tibia, a few scattered on distal portion of femur; femur-tibia articulation functional; tarsal and claw digitules distinctly expanded at apices. Anal opening 65–84 µm wide, on ventral body surface, surrounded by sclerotic anal ring 85–104 µm wide bearing 14–18 setae, base of each seta surrounded by small pores.

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.

Holotype of Opisthoscelis thurgoona (here designated): AUSTRALIA: New South Wales: 1 adult female (3.4 mm long, 2.8 mm wide):ex leaf gall, Eucalyptus melliodora, 2 km NE of Thurgoona, Ettamogah Road, -36.03°; 146.98°, 8 Feb., 2004, PJG (ANIC).

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, Eucalyptus melliodora, 1 km N of Thurgoona, Table Top Road, -36.05°; 146.98°, 30 Dec., 1991, PJG (ANIC, except 1 adult female in BMNH, 1 adult female in USNM).

Adult females of Opisthoscelis thurgoona are unusual amongst Opisthoscelis and Tanyscelis species in having each leg well developed, and 7 clearly separate antennal segments. These features, in combination with the ventral anal opening, with a sclerotic anal ring bearing numerous setae, each of which is surrounded at the base by a number of minute pores, give the adult female of Opisthoscelis thurgoona the resemblance of a species of Lachnodius. Morphological clues of the closer relationship between Opisthoscelis thurgoona and the other Opisthoscelis species (recovered by analysis of DNA sequence data (NBH, unpublished data)) are the presence of macrotubular duct clusters on the dorsum (possibly homologous to the cribriform plates found in Opisthoscelis subrotunda and Opisthoscelis beardlseyi) and only 4 fleshy setae on each antenna (5 on species of Lachnodius). Opisthoscelis thurgoona is known only from a single roadside location, 1 km north of Thurgoona near Albury in New South Wales. Opisthoscelis thurgoona is most similar to Opisthoscelis tuberculata (described as new below) and the two species are compared under the comments for the latter.

Live eggs of Opisthoscelis thurgoona are laid in the gall cavity and are pink to orange in colour, whereas newly hatched first-instar nymphs are orange. The chromosome number is 2n = 18 (LGC, unpublished data).

The species name is taken from the type locality. It is a noun in apposition.

(Fig. 1k). Female. On leaf. Pit-shaped gall, opening round to oblong, 2.0–3.0 mm wide, surrounded by raised lip, ca 0.2 mm high, apparently on adaxial leaf surface; gall hemispherical on side of leaf opposite opening, 1.0–1.5 mm high.

Male. Not known.

(Fig. 9) (n = 4). Body broadly oval in outline, length 2.0–4.0 mm, greatest width 1.4–3.2, venter larger than dorsum. Eyes each 40–50 um wide, slightly ventral of line of marginal setal fringe. Antennae 7-segmented, moderately long (0.72–1.10 mm) and slender; with segment III longest (180–290 µm) and segment VII shortest (70–95 µm); segment I with 3 hair-like setae; II with 7–11 hair-like setae; III with 8–12 hair-like setae; IV 5–7 hair-like setae; V with 4–6 hair-like setae + 1 fleshy seta; VI with 3 or 4 hair-like setae + 1 fleshy seta; VII with ca 6 hair-like setae + 3 fleshy setae. Frontal lobes difficult to discern, each perhaps 200–280 µm long, 110–120 µm wide. Tentorial box 125–200 mm long; labium 100–150 mm long, 120–140 mm wide, 2-segmented with basal segment indicated by presence of setae; pump chamber 20–27 µm long, 17–20 µm wide. Spiracles 90–140 mm long, 55–95 mm wide across atrium. All legs well developed, hind legs larger than fore or mid legs: fore leg with coxa 240–400 µm long, trochanter + femur 370–550 µm long, tibia 280–440 µm long, tarsus 110–160 µm long; mid leg with coxa 270–450 µm long, trochanter + femur 380–580 µm long, tibia 280–460 µm long, tarsus 110–150 µm long; hind leg with coxa 320–520 µm long, trochanter + femur 400–620 µm long, tibia straight, 310–500 µm long, tarsus 120–180 µm long; each leg with claw 35–45 µm long; small translucent pores scattered on both surfaces of all hind-leg segments, densest on tibia; femur-tibia articulation functional; tarsal and claw digitules distinctly expanded at apices. Anal opening 53–75 µm wide, on ventral body surface, surrounded by sclerotic anal ring 68–100 µm wide bearing 12–14 setae, each 150–175 µm long with setal base surrounded by 4–10 minute pores.

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.

Holotype of Opisthoscelis tuberculata (here designated): AUSTRALIA: New South Wales: 1 adult female (4.0 mm long, 3.2 mm wide):ex open top pit gall in leaf midrib, Eucalyptus sp., Penrith, 24 Nov., 1899, WWF No. 304 (ASCU).

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 Eucalyptus sp. (perhaps Eucalyptus microcarpa), ca 10 km NNW of Benalla, Midland Highway, near turnoff to Hwy C371 to Tocumwal, -36.47°; 145.94°, 29 Dec., 2008, PJG, NH205 (ANIC).

Adult females of Opisthoscelis tuberculata are closely related to Opisthoscelis thurgoona with which they share well-developed fore, mid and hind legs, 7 distinct antennal segments, a ventral anal opening, with a sclerotic anal ring bearing numerous setae, each of which is surrounded at the base by a number of minute pores. These features of the adult females of Opisthoscelis tuberculata and Opisthoscelis thurgoona are shared with species of Lachnodius. One morphological clue of the closer relationship between Opisthoscelis tuberculata and Opisthoscelis thurgoona and the other Opisthoscelis species (recovered by analysis of DNA sequence data (NBH, unpublished data)) is the presence of macrotubular duct clusters on the dorsum (possibly homologous to the cribriform plates found in Opisthoscelis subrotunda and Opisthoscelis beardsleyi). However, as in Lachnodius, Opisthoscelis tuberculata has 5 fleshy setae on each antenna (4 on Opisthoscelis thurgoona).

The adult female of Opisthoscelis tuberculata can be distinguished readily from that of Opisthoscelis thurgoona as follows (features of Opisthoscelis thurgoona in parentheses): (1) 5 fleshy setae on each antenna (4 on each); (2) dorsal ducts in clusters of up to 30 ducts (2–8 ducts per cluster); and (3) the adult female sits in an open-top leaf pit (in an enclosed leaf gall in Opisthoscelis thurgoona).

The holotype of Opisthoscelis tuberculata is from the collection of W.W. Froggatt and the female was removed from an open pit gall in a leaf midrib and slide-mounted by J.W. Beardsley. Froggatt’s first accession notebook (Gullan 1984b) records collection #304 as “Dactylopius eucalypti.? Penrith (no.2) (Berlese 234). small ascelis like gall”. The reference to Berlese may refer to Professor Antonio Berlese (dal Laboratorio di entomologia agraria, Portici, Italy), because two nearby entries in the notebook say “Sent to Berlese” and give different numbers. The 3 adult females from Victoria were collected dead (post-oviposition), in association with their salmon-pink eggs, under loose bark just above the ground on the main trunk of a stunted tree (about 1.5 m high) growing in the middle of a picnic area car park. Several leaves on the tree had vacated pit galls (Fig. 1k), which are presumed to have been the feeding sites of the females prior to their movement to the bark for oviposition. The live eggs were maintained in a small plastic container and started eclosing on 1 Jan., 2009; DNA was extracted from the first-instar nymphs by NBH.

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 Lachnodius. We assume that his name refers to the sclerotised dorsal duct clusters.

(Fig. 1l). Female. On leaf. Raised, rather nipple-like excrescences; height ca 2 mm above leaf and projecting 1 mm below, width 3–5 mm. Gall opening star-like, with fissures radiating from central orifice, apparently on adaxial leaf surface.

Male. Unknown.

(Fig. 10) (n = 7). Body outline circular, with venter broader than dorsum of mature specimens; length 3.0–3.9 mm, greatest width 2.8–3.7 mm; abdomen about as long as head + thorax. Eyespots on dorsum near margin, each 48–58 mm wide. Antennae 3–5 segmented, each 185–230 mm long. Frontal lobes difficult to discern in some specimens, each 330–510 µm long, 185–230 µm wide. Tentorial box 480–530 mm long. Labium 140–155 mm long, 125–175 mm wide. Pump chamber 28–33 µm long, 35–38 µm wide. Spiracles 135–170 mm long, 65–90 mm wide across atrium. Fore and mid legs reduced but segmented, 245–410 µm long. Hind legs slender and elongate; coxa on a fleshy projection, coxa 650–745 µm long, trochanter + femur 820–880 µm long, tibia straight, 645–680 µm long, tarsus 350–410 µm long; claw and digitules present but reduced; translucent pores present on both surfaces of each leg segment except trochanter; each side of trochanter with 3 campaniform sensilla; femur-tibia articulation functional. Anal opening on venter, 58–68 µm wide, anal ring 83–110 µm wide, with 14–20 setae.

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.

Holotype of Opisthoscelis ungulifinis (here designated): AUSTRALIA: South Australia: 1 adult female (3.9 mm long, 3.7 mm wide): ex gall, Eucalyptus ?oleosa, 2 km SW of Oodla Wirra, 5 Dec., 1981, PJG (ANIC).

Paratypes: AUSTRALIA: South Australia: 6 adult females, 24 first-instar nymphs: same data as holotype (ANIC).

Adult females of Opisthoscelis ungulifinis can be recognised easily by the 1 or 2 blunt spines on each of the marginal projections on the posterior abdominal segments, and by having the hind coxae each set on a fleshy projection. Adult females of Opisthoscelis ungulifinis are also unique amongst Opisthoscelis species in having microtubular ducts. The adult females are most similar to those of Opisthoscelis serrata, which also have marginal projections each with 1 or 2 spines; however these spines are conical in Opisthoscelis serrata. Adult females of Opisthoscelis ungulifinis can be further separated from those of Opisthoscelis serrata by lacking marginal projections cephalad of abdominal segment IV (present on all abdominal segments of Opisthoscelis serrata), and by having a marginal fringe of elongate setae (absent in Opisthoscelis serrata).

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 Schrader (1863: 7) as follows: “In another species Opisthoscelis gracilis, which I have observed, the oviparous female is rather slender, and the legs are still longer and thinner than in the species before noticed [Opisthoscelis subrotunda], and the male has no anal setae.” There is no illustration and Schrader’s specimens were destroyed in WWII (Gullan 1984a). Froggatt (1894a) considered Opisthoscelis gracilis to be a synonym of Opisthoscelis subrotunda. However, Schrader’s description of the adult female having long thin legs and a slender body, and the adult male with no anal setae, suggest that this was a species of Tanyscelis. In the absence of type specimens, it is impossible to determine which species this name applies to and we here consider it to be a nomen dubium.

Fuller’s (1897: 1346) original description of Opisthoscelis conica is very brief: “Upon one side of the leaf arises the conical apex, whilst on the other the gall protrudes as a hemisphere.” Later Fuller (1899) published a more detailed description with an illustration of three galls of females on a leaf and a hind leg of an adult female (plate XV, figs 33–34). The galls in Fuller’s drawing are very similar to syntype galls housed in the SAMA (see “Material examined” for details), but no insects have been found in the SAMA or elsewhere (e.g., not in the South African National Collection of Insects where some other Fuller material is housed). The type locality of Opisthoscelis conica was given as Midland Junction in Fuller (1897) but as Swan River in Fuller (1899), however it is unlikely that there were two localities for this species because Fuller published the second paper from South Africa based on notes that he had made during eight or nine months spent collecting in Perth prior to his 1897 paper. Midland Junction, now the suburb of Midland, is several km east of the Swan River and, in the 1890s, it was a railway station and associated village.

(Fig. 2a, b). Female. On leaf. Height 5.0–7.0 mm (= total height above and below leaf blade), width 5.0–9.5 mm. Gall opening slit-like, 0.5–0.8 mm long; on abaxial or adaxial surface, but all galls opening on same surface on any one leaf. Side of gall with opening conical, other side convex, globose (Fig. 2b).

Male. Not known.

(Fig. 11) (n = 8). Body outline turbinate, margin incised at intersegmental boundaries, length 2.5–3.7 mm, greatest width 1.1–2.5 mm; abdomen tapered, about as long as head + thorax, extending far beyond femur. Eyespots small, each 18–25 mm wide. Antennae 1-segmented, each 35–108 mm long. Frontal lobes difficult to see, each ca 170 µm long, 200 µm wide. Tentorial box 350–470 mm long. Pump chamber 30–43 µm long, 38–48 µm wide. Labium 85–113 mm long, 95–140 mm wide. Spiracles 70–105 mm long, 45–63 mm wide across atrium. Fore and mid legs small sclerotic protuberances; fore leg 20–35 µm long, with 6–8 setae; mid leg larger, 38–80 µm long, with 10–12 setae. Hind legs robust and elongate; coxa 180–260 µm long, trochanter about as a long as femur, the two combined 450–610 µm long, tibia and tarsus fused, forming straight, sword-like segment 720–1880 µm long; claw and digitules absent; translucent pores dense throughout distal region of dorsal and ventral surfaces of tibiotarsus; femur-tibia articulation poorly developed. Anal opening poorly formed, 15–28 µm wide, surrounded by dense cluster of setae.

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: “Opisthoscelis Schrader / conica, Fuller, sp. n. Type / Perth, W. Australia / C. Fuller / 28.7.97” and “ Perth, W. A. / Opisthoscelis / conica Fuller / m/s / 28.7.97” (SAMA).

The adult female of Tanyscelis conica would be difficult to confuse with the adult female of any other known species of Tanyscelis. The fusion of the hind tibia and tarsus into a broad, sword-like appendage is unique. Both the form of the dorsal setae (robust and slightly curved) and their dense distribution also separate adult females of Tanyscelis conica from those of other species. Known populations of Tanyscelis conica occur in two disjunct clusters, one centered around Perth in southwest Western Australia, and the other in forest reserves in northwestern Victoria and southeastern Australia (e.g., Danggali Conservation Park, Wyperfield National Park, Hattah-Kulkyne National Park). Despite the great distance separating the two, ca 2000 km, the morphology is homogeneous. Tanyscelis conica is the only species of Tanyscelis known from Western Australia. Only one species of Opisthoscelis, the type species Opisthoscelis subrotunda, has been collected from Western Australia, in the Kimberley region, which is a great distance (> 1, 500 km) north of Perth. Tanyscelis conica is also the only known species of Tanyscelis or Opisthoscelis known to feed on mallee eucalypt species in the section Dumaria, and along with Opisthoscelis ungulifinis, is one of only two species in this group known to feed on eucalypts in the section Bisectae.

We have examined all available type specimens of Froggatt’s Opisthoscelis convexa and Opisthoscelis globosa (the latter now Opisthoscelis ruebsaameni) (see under “Material examined” for details of types) and inasmuch as the poor quality of the specimens allows comparison, we consider these adult females to be identical. Froggatt (1929) described these two species on the same page of his paper, with Opisthoscelis convexa described first, and based his new species on specimens of each from both Victoria (Diamond Creek for Opisthoscelis convexa but not specified for Opisthoscelis globosa Froggatt) and from New South Wales (Gosford, Macksville and Warrah for Opisthoscelis convexa and Hornsby for Opisthoscelis globosa Froggatt). He stated that the gall of the female of his Opisthoscelis globosa was larger and more swollen, and the adult female was broader with more irregular transverse bands of spines on the dorsal abdominal segments than on the female of Opisthoscelis convexa. We did not observe a striking difference in the dorsal abdominal spines beween Froggatt’s specimens of Opisthoscelis globosa and Opisthoscelis convexa. The size differences he mentions probably relate to the age of the females.

(Fig. 2c, d). Female (Fig. 2c). On stem and base of petiole. Height 3.0–7.7 mm, width 4.5–8.6 mm, length of basal attachment 4.6–11.8 mm. Gall opening slit-like to oblong, 0.1–0.4 mm wide, 0.4–1.7 mm long (paralectotype galls with slit-like opening mostly 0.1 x 1.2 mm). Gall globose, surface variable, distal area frequently with 1 or 2 concentric circular scars, distal surface above scars smooth and lighter in colour than surrounding stem tissue.

Male (Fig. 2d). On either leaf surface, occasionally on petiole or stems. Height 1.4–3.7 mm, width 1.0–1.6 mm, length of basal attachment 1.3–2.6 mm. Gall cylindrical to conical, distal margin dentate, opening slit-like to round, 0.5–1.7 mm wide, opposite side of leaf swollen.

(Fig. 12) (n = 24). Body outline broadly turbinate, outline of head + thorax + abdominal segment I entire, abdominal margin weakly incised between each segment, length 2.3–4.4 mm, greatest width 1.9–5.0 mm; abdomen tapered, about as long as head + thorax. Eyespots each 25–83 mm wide, on dorsum among or medial to marginal spines. Antennae 1-segmented, in form of low convex plate or knob; width 25–78 mm. Frontal lobes difficult to see, each ca 220 µm long, 230 µm wide. Tentorial box 350–600 mm long. Pump chamber 50–63 µm long, 45–55 µm wide. Labium 73–160 mm long, 80–125 mm wide. Spiracles 100–205 mm long, 48–118 mm wide across atrium. Fore and mid legs small sclerotic protuberances, 20–45 µm long. Hind legs slender and elongate; coxa 360–670 µm long, hirsute, trochanter + femur 480–880 µm long, tibia 1000–1610 µm long, tarsus 75–250 µm long; translucent pores dense on both dorsal and ventral surfaces of tibia and tarsus; femur-tibia articulation non-functional, tibia fixed in orientation parallel to long axis of femur; claw and digitules absent. Anal opening poorly formed, 8–20 µm wide; sclerotic anal ring and anal ring setae absent.

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.

Lectotype of Opisthoscelis convexa Froggatt (here designated): AUSTRALIA: Victoria: 1 adult female on WWF original slide-mount with another female, lectotype (3.4 mm long, 4.0 mm wide) closest to middle of slide, with labels: “Opisthoscelis / convexa / n sp / Eucalyptus / NSW & Vic.” and “O. convexa” (ANIC).

Paralectotypes of Opisthoscelis convexa: AUSTRALIA: Victoria: 1 adult female on same slide-mount as lectotype: same data as lectotype (ANIC); 1 adult female, WWF original slide-mount labelled: “Opisthoscelis / convexa / n sp / N. S. Wales / WWF” and “Finished O. convexa” (ANIC); 2 adult females plus associated dry galls on stem, 6 slides of embryos and first-instar nymphs (several hundred) and 7 other dry galls of females: ex Eucalyptus macrorrhyncha, Diamond Creek, Oct., 1921, C. French, 1055, ASCT00004874 and ASCT00004875 (ASCU).

Lectotype of Opisthoscelis globosa Froggatt (here designated): AUSTRALIA: New South Wales: 1 adult female (3.8 mm long, 4.0 mm wide): WWF original slide-mount labelled: “Opisthoscelis / globosa / n sp / N. S. Wales / WWF” and “O. globosa” (ANIC).

Paralectotype of Opisthoscelis globosa: 1 adult female: WWF original slide-mount labelled: “Opisthoscelis / globosa / n sp” and “O. globosa” (ANIC).

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 Opisthoscelis convexa, the collection data for these specimens are not listed in Froggatt’s (1929) description of the species and thus we do not consider them to be part of the type material.

Adult females of Tanyscelis convexa can be recognised easily by the large dorsal spines found on the dorsomedial areas of the thorax and abdomen and in a ventral submarginal row extending from the anterior margin of the head to the posterior spiracles. Tanyscelis convexa is also the only species of Tanyscelis to have no tubular ducts or multilocular pores on the dorsum. DNA sequence data support a sister relationship between Tanyscelis convexa and Tanyscelis maculata + Tanyscelis maskelli (NBH, unpublished data). Tanyscelis convexa is one of the more frequently collected, and broadly distributed, species of Tanyscelis. It is known to occur along the coast from Adelaide in South Australia to Cairns in north Queensland. Adult females from near Swan Hill but in New South Wales, and from Canberra differ from all other collections of Tanyscelis convexa in having the ventromarginal band of spines poorly developed. Samples from Benalla, Shepparton, and Nagambie (more or less between but about 100 km farther south than the Canberra and Swan Hill samples) have the ventromarginal band of spines well developed. Tanyscelis convexa has been collected exclusively from eucalypt species in the section Adnataria.

(Fig. 2e). Female. On leaf. Thorn-like, height 2.2–3.0 mm, width 3.0–4.8 mm, length of basal attachment 5.0–6.0 mm. Gall opening slit-like, 0.5–0.8 mm long; on adaxial leaf surface for several galls examined. Base broad and globose with circular scar; apex tapering to blunt point. Live adult females of Tanyscelis grallator have white powdery wax in a band across dorsum of each body segment, with intersegmental areas bare of wax. The female is small in relation to the size of the gall cavity and eggs are laid into the cavity, which can become filled.

Male. Not known.

(Fig. 13) (n = 10). Body turbinate, head, thorax, and anterior abdominal segments with intersegmental boundaries indistinct along margin, intersegmental boundaries incised along posterior abdominal margin; body length 2.4–2.7 mm, greatest width 1.7–2.0 mm; abdomen tapered, about as long as head + thorax, not extending beyond outstretched femur. Large (115–215 mm long), papilliform fleshy protuberance on each side of head in place of eyes; apex of each protuberance with differentiated area, sometimes surrounded by invagination. Antennae each 1-segmented, 55–88 mm long. Frontal lobes difficult to see, each 125–255 µm long, 145–245 µm wide. Tentorial box 310–410 mm long. Labium 88–113 mm long, 68–93 mm wide. Pump chamber 30–38 µm long, 30–38 µm wide. Spiracles 83–128 mm long, 53–73 mm wide across atrium. Fore leg 43–88 µm long; mid leg 68–163 µm long. Hind leg slender and elongate; coxa cylindrical, 520–630 µm long, trochanter + femur 770–960 µm long, tibia 1200–1900 µm long, tarsus 255–420 µm long; translucent pores dense on both surfaces of tibia, few on tarsus; trochanter with 2 campaniform sensilla on each side; femur-tibia articulation functional; claw and digitules present but reduced. Anal opening 10–20 µm wide, without distinct sclerotic anal ring.

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.

Holotype (here designated): AUSTRALIA: Queensland: 1 adult female (2.4 mm long, 2.0 mm wide): ex conical gall on leaf, Eucalyptus sp., 29 km N of Normanton, -17.47°; 141.18°, 1 Oct., 2006, LGC, LGC00666 (ANIC).

Paratypes: AUSTRALIA: Queensland: 2 adult females: ex leaf gall, Eucalyptus sp. (ironbark), 13 km N of Injune, 7 Dec., 1993, LGC (ANIC); 1 adult female: ex Eucalyptus sp., Charters Towers, Oct., 1955, ARB, INSECOLL 0-067153 (QDPI); 1 adult female: ex Eucalyptus crebra, Clermont, Nov., 1938, INSECOLL 0-067159 (QDPI); 3 adult females: ex leaf galls, Eucalyptus sp. (ironbark), Dunmore State Forest, 2 May, 1995, G. Harper (ANIC); 4 adult females: ex leaves, Eucalyptus crebra, Emu Vale, 15 Mar., 1939, INSECOLL 0-067161, 0-067162 (QDPI); 1 adult female: ex gall, Eucalyptus crebra, Mt Walsh Nat. Park, picnic ground, 267 m, -25.57°; 152.05°, 11 Oct., 2003, LGC and M. D. Crisp, LGC00029 (ANIC); 1 adult female: no data, INSECOLL 0-067168 (QDPI).

Adult females of Tanyscelis grallator are most similar to those of Tanyscelis mollicornuta. Most distinctively, adult females of both species have fleshy protuberances in place of eyes. These protuberances are much longer on Tanyscelis mollicornuta (≥ length of hind tarsus) than on Tanyscelis grallator (<< length of hind tarsus), which has additional protuberances along the dorsal midline plus the submargin of the thorax. Each protuberance on the head of Tanyscelis grallator has the apex much more differentiated than the similar protuberances of Tanyscelis mollicornuta, and it seems possible that this area is light sensitive. Adult females of Tanyscelis grallator can be distinguished further from those of Tanyscelis mollicornuta by having much longer hind legs, each with a cylindrical coxa (more conical in Tanyscelis mollicornuta), and the ratio of the hind trochanter + femur to tibia + tarsus << 1 (ca 1 in Tanyscelis mollicornuta). Tanyscelis mollicornuta is known from only one location, in Queensland, east of Western Creek State Forest, near Milmerran. Samples of Tanyscelis grallator have been collected from five locations in Queensland, including a collection from the eastern edge of Kumbarilla State Forest, < 50 km to the north of Milmerran. Tanyscelis mollicornuta and Tanyscelis grallator have only been found on eucalypts in the subgenus Adnataria.

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 Opisthoscelis recurva with Opisthoscelis maculata. We could find no morphological differences between the adult females of each, although type adult females of Opisthoscelis recurva tend to be larger than those of Opisthoscelis maculata. Furthermore, the ‘recurved’ gall morphology from which the name of the junior synonym was derived is found occasionally in galls of smaller specimens that match the types of Opisthoscelis maculata. The type locality for Opisthoscelis maculata is Bendigo, Victoria, with galls reported from Eucalyptus leucoxylon and Eucalyptus gracilis (Froggatt 1894b), whereas Opisthoscelis recurva was described from specimens collected on the hills at Warrah Station, a farming property near Willow Tree [not Willowtree as in the description], New South Wales, on an undetermined species of Eucalyptus (Froggatt 1929).

(Fig. 2f, g). Female. On stem (mostly), fruit, leaf petiole and midrib. Conical (Fig. 2f) to thorn-like (Fig. 2g), height 3.2–16.2 mm, width 2.7–6.9 mm, length of basal attachment 3.6–11.2 mm. Gall opening oblong, 0.1–0.4 mm wide, 0.3–1.2 mm long. Base of gall broad, surface smooth, distal portion attenuated and annulated, may be recurved, 1–6 rings.

Male. On stem and either leaf surface (Fig. 2f), height 1.9–8.0 mm, width 0.9–2.8 mm, length of basal attachment 1.1–3.8 mm. Gall thin-walled and cylindrical, distal edge crenulate, opening round to oblong, 0.3–2.4 mm maximum wide.

(Fig. 14) (n = 53). Body outline ovate to turbinate, length 1.7–5.6 mm, greatest width 0.9–4.6 mm; abdomen tapered, quite small relative to thorax and head. Eyespots large, each 45–123 mm wide, highly convex, surrounded by dense cluster of stout setae. Antennae each 1-segmented, 45–520 mm long. Frontal lobes difficult to see, each 180–650 µm long, 180–580 µm wide. Tentorial box 260–520 mm long. Pump chamber 33–60 µm long, 34–55 µm wide. Labium 50–163 mm long, 60–138 mm wide. Spiracles 85–240 mm long, 38–130 mm wide across atrium. Fore and mid legs small sclerotic protuberances, 15–120 µm long. Hind legs slender and elongate; coxa 240–730 µm long, trochanter + femur 240–930 µm long, tibia 840–4500 µm long, tarsus 140–350 µm long; translucent pores sparsely scattered on dorsal and ventral surfaces of tibiotarsus; trochanter with 3 or 4 campaniform sensilla on each side; femur-tibia articulation non-functional, tibia fixed in orientation parallel to long axis of femur; claw and digitules absent. Anal opening poorly formed, 10–45 µm wide, sclerotic anal ring and anal ring setae absent.

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.

Lectotype of Opisthoscelis maculata (here designated): AUSTRALIA: Victoria: 1 adult female (2.3 mm long, 2.2 mm wide): ex dry gall on stem, with printed label: “No. 1787 E / GALL MAKING COCCIDS. / Opisthoscelis maculata, Frogtt. / Female galls on Eucalyptus / Bendigo, Victoria“, ASCT00004880 (ASCU).

Paralectotypes of Opisthoscelis maculata: AUSTRALIA: Victoria: 1 adult female, same data as lectotype (ASCU).

Lectotype of Opisthoscelis recurva (here designated): AUSTRALIA: New South Wales: 1 adult female (5.2 mm long, 4.1 mm wide): on WWF original slide-mount labelled: “Opisthoscelis / recurva / n sp / Warrah NSW” (ANIC).

Paralectotypes of Opisthoscelis recurva: AUSTRALIA: New South Wales: 1 adult female and 2 dry galls of female on base of leaves (female ex one of these galls slide-mounted by PJG 2008), with WWF labels: “Warrah / Sydney, N.S.W. / W. W. Froggatt. / Eucalypt / 8.5.1921” and “1045”, ASCT00004859 & ASCT00004860 (ASCU); 1 adult female and 2 slides of first-instar nymphs ex dry gall on leaf, plus other associated dry galls, with 2 labels: “1579 / Opisthoscelis / recurva / n sp MS / WWF Warrah, N.S.W.” and “Opisth. recurva. (Frogg. MS) / on Eucalyptus sp. / Warrah. N.S.W., Australia” (BMNH). Note: The labels on the 2 dry type galls of Opisthoscelis recurva at ASCU say “Warrah / Sydney” but this is an error; Warrah is property near Willow Tree, in northeastern N.S.W. The paralectotypes in the BMNH are labelled correctly.

Adult females of Tanyscelis maculata are very similar to those of Tanyscelis maskelli. The latter can be distinguished by the much smaller eyes, the distinct marginal fringe of setae, and the longer dorsal setae on the abdomen and along the margin. Tanyscelis maculata is known from many more collections, and appears to be common in the south of Australia, whereas Tanyscelis maskelli has a more northern distribution. Populations of both species have been collected in New South Wales, although never from the same site. Except for one sample of Tanyscelis maculata collected by J. W. Beardsley, apparently from Eucalyptus macrorrhyncha (subgenus Eucalyptus: section Aromatica), and Froggatt’s (1894b) unconfirmed record from Eucalyptus gracilis (subgenus Symphyomyrtus: section Bisectae), all material of Tanyscelis maskelli and Tanyscelis maculata has been collected from eucalypt species in the section Adnataria (subgenus Symphyomyrtus).

In the original description, Froggatt (1894b) states that he collected this species at Cooma, Maitland, Newcastle, and a dozen localities within a radius of 20 miles [32 km] of Sydney, but that a constant locality was Flemington on Eucalyptus siderophloia, the large-leaved box. Flemington is in western Sydney and now in the suburb of Homebush West Subsequently, Froggatt (1929) treated Flemington as the type locality, as did Miller and Gimpel (2000) in their catalogue of the Eriococcidae.

(Fig. 2h, i). Female (Fig. 2h). On stem. Conical (Fig. 2h) to thorn-like, height 6.4–14.1 mm, width 3.7–7.8 mm, length of basal attachment 5.2–11.0 mm. Gall opening round to oblong, 0.2–1.8 mm wide. Base of gall broad, surface rugose, distal portion attenuated and annulated, with 2–7 rings.

Male (Fig. 2i). On stem and either leaf surface. Tubular, narrowing slightly towards apex, height 2.1–7.0 mm, width 1.0–2.1 mm, length of basal attachment 1.3–4.7 mm. Gall opening indistinct to round, 0.3–1.7 mm wide. Gall thin-walled and cylindrical, distal edge surrounding opening crenulate.

(Fig. 15) (n = 16). Body outline ovate to turbinate, length 3.5–6.2 mm, greatest width 3.1–5.1 mm; abdomen tapered, quite small relative to thorax and head. Eyespots each 20–45 mm wide, dorsad of margin fringe setae. Antennae 1-segmented, 43–130 mm long. Frontal lobes difficult to see, each 300–350 µm long, 350–520 µm wide. Tentorial box 440–1400 mm long. Labium 125–160 mm long, 83–150 mm wide. Spiracles 120–310 mm long, 60–120 mm wide across atrium. Fore and mid legs small sclerotic protuberances, 25–55 µm long. Hind legs slender and elongate; coxa 500–1170 µm, trochanter + femur 510–1222 µm, tibia 1820–2730 µm long, tarsus 115–360 µm; translucent pores on both surfaces of tibia and tarsus; trochanter with 3 or 4 campaniform sensilla on each side; femur-tibia articulation non-functional, tibia fixed in orientation parallel to long axis of femur; claw and digitules absent. Anal opening poorly formed, 15–35 µm wide, sclerotic anal ring and anal ring setae absent.

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.

Lectotype of Opisthoscelis maskelli (here designated): AUSTRALIA: New South Wales: 1 adult female (4.02 mm long, 3.24 mm wide): ex dry gall on stem, with printed label: “No. 1944 E / GALL MAKING COCCIDS. / Ophisthoscelis[sic] maskelli, Frogtt. / Male and female galls on Eucalyptus / Flemington, N.S.W.” ASCT00004851 (ASCU).

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 / Opisthoscelis / MASKELLI / n. sp. / female / 1891 Froggatt W.M.M” and “Co Type / A gall making scale / insect, showing /the long hind / legs. / B.M. 1945.121” (BMNH); 2 adult females (slide-mounted by PJG from dry collection): ex dry stem galls with labels: “Opisthoscelis maskelli / Frogtt / on Eucalyptus siderophloia / Sydney – Australia / ex coll. W.W. Froggatt” and “Opisth Maskelli / ex Coll W. Maskell.” (BMNH).

Adult females of Tanyscelis maskelli are very similar to those of Tanyscelis maculata [see comments under Tanyscelis maculata].

(Fig. 3a). Female. On stem. A rounded swelling; gall opening slit-like; base of gall broad, gall surface striated, distal part a narrow truncate cone bearing orifice.

Male. On stem; similar to gall of female but without apical cone; gall opening irregularly shaped.

(Figure 16) (n = 19). Body rotund, with 3 or 4 very large dorsal humps, body margin ovate, length 1.4–3.3 mm, greatest width 0.9–ca 3 mm; abdomen appears to be tapered (truncate in all slide-preparations, but clearly tapered in photograph of adult female in life). Eyes each 60–85 µm wide, convex, with base not parallel-sided or perpedicular to body surface. Antennae 1-segmented, 95–125 mm long. Frontal lobes often difficult to detect, each ca 150 µm long, 190 µm wide. Tentorial box 325–450 mm long. Labium 115–150 mm long, 100–110 mm wide. Pump chamber 30–38 µm long, 35–40 µm wide. Spiracles 125–170 mm long, 55–100 mm wide across atrium. Fore and mid legs small sclerotic stubs, 18–45 µm long. Hind leg slender and elongate; coxa 370–430 µm long, trochanter + femur 480–550 µm long, tibia straight, 750–1000 µm long, tarsus 410–540 µm long; translucent pores scattered over both surfaces tibia, plus a few on femur and tarsus; femur-tibia articulation non-functional, base of tibia fixed at obtuse angle to femur; each side of hind trochanter with 3–6 campaniform sensilla; claw and digitules absent. Anal opening 8–20 µm wide, anal ring poorly developed, with ca 6 setae.

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.

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, Eucalyptus microcarpa [original label incorrectly has Eucalyptus odorata; see Comments below], Aldinga Beach, 2 Oct., 1965, H. M. Brookes, Specimen Index No. 44/65 (ANIC).

Paratypes: AUSTRALIA: South Australia: 2 adult females, on same slide as holotype (ANIC); 6 adult females: ex stem galls, Eucalyptus microcarpa, Aldinga Beach, 9 Jan., 1964, H. M. Brookes, Specimen Index No. 4/64 (ANIC); 10 adult females: same data except 2 Oct., 1965, Specimen Index No. 44/65 (ANIC); 4 adult females: ex galls, Eucalyptus sp., Aldinga Beach, 11 Mar., 1966, HMB (ANIC); 7 adult females: ex galls, Eucalyptus sp., 1.5 km ESE of Moorlands, 6 May, 1980, PJG (ANIC).

Adult females of Tanyscelis megagibba are most similar to those of Tanyscelis villosigibba, in the sense that the dorsal surface of both species is dominated by a series of large humps. These are most spectacular in mature females of Tanyscelis megagibba, but lack the dense covering of setae, each born on a raised fleshy base, that is characteristic of the adult females of Tanyscelis villosigibba. Adult females of Tanyscelis villosigibba are also distinctive in having highly convex eyes, with the base of each eye parallel-sided and perpendicular to the body surface, whereas in Tanyscelis megagibba the base of each eye is not parallel-sided. Tanyscelis megagibba is known from only two localities, ca 100 km apart, in South Australia, whereas Tanyscelis villosigibba is known from four localities in Queensland.

The original labels on the slides from two of the three collections of Tanyscelis megagibba made by HMB have the host as Eucalyptus odorata. However, her subsequent notes specify that this was a misidentification and the collections were made from Eucalyptus microcarpa (not Eucalyptus odorata).

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.

(Fig. 17) (n = 4). Body turbinate, margin incised at intersegmental boundaries, length 2.0–2.6 mm, greatest width 1.4–1.5 mm; abdomen tapered, about as long as head + thorax, extending beyond femur. Eyespot undetected; large (330–425 µm long), papilliform fleshy protuberance on each side of head in place of eyes. Antennae 1-segmented, 38–123 mm long. Frontal lobes difficult to see, each ca 120 µm long, 110 µm wide. Tentorial box 225–315 mm long. Labium 100–113 mm long, 60–75 mm wide. Pump chamber 28–33 µm long, 28–33 µm wide. Spiracles 85–120 mm long, 43–50 mm wide across atrium. Fore leg 50–65 µm long; mid leg 58–105 µm long. Hind leg slender and elongate; coxa conical, 300–430 µm long, trochanter + femur 580–610 µm long, tibia 630–720 µm long, tarsus 290–300 µm long; translucent pores on proximal-lateral area of each coxa, plus on both surfaces of tibia and tarsus; trochanter with 3 campaniform sensilla on each side; femur-tibia articulation functional; claw and digitules present but reduced. Anal opening ventral, 13–15 µm wide, without distinct sclerotic anal ring, surrounded by rugose area of cuticle bearing ca 10 setae.

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.

Holotype (here designated): AUSTRALIA: Queensland: 1 adult female (2.0 mm long, 1.5 mm wide): ex gall on leaf, Eucalyptus populnea, 3 km WSW of Millmerran, Turallin Rd, -27.88°; 151.24°, 1 May, 1995, LGC (ANIC).

Paratypes: AUSTRALIA: Queensland: 5 adult female: same data as holotype (ANIC).

Adult females of Tanyscelis mollicornuta are most similar to those of Tanyscelis grallator [see comments under Tanyscelis grallator].

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 Froggatt (1894b) described Opisthoscelis pisiformis, he listed three host species of eucalypts – Eucalyptus melliodora, Eucalyptus piperita and Eucalyptus resinifera – from three different localities in New South Wales –Bathurst, Thornleigh and Sutherland. Even though we designate a lectotype below, the type locality of this species continues to encompass these three localities because the collection locality of the lectotype is not known. Although Miller and Gimpel (2000) listed the BMNH as holding syntypes, the data associated with the four BMNH collections of dry galls labelled as Opisthoscelis pisiformis either clearly show that the material was collected subsequent to description of this species or the data are inadequate to determine type status. Probably only two of the four BMNH collections belong to this species. Insects extracted from dry galls of one collection were slide-mounted by PJG in 1984 and are listed below in Material examined.

(Fig. 3b, c). Female (Fig. 3a). On leaf. Height 4.0–9.6 mm, width 4.0–6.4 mm, length of basal attachment 2.9–8.9 mm. Gall opening an almost closed slit in young galls, in mature galls slit-like to oblong, 0.1–1.8 mm wide, 0.6–0.8 mm long; on abaxial (lower) leaf surface. Gall opening on small raised or low conical protrusion, opposite side of leaf sub-spherical, basal attachment may be constricted; surface green, sometimes with reddish tinge, leaf glands enlarged.

Male (Fig. 3b). On stem and leaf (probably either surface), height 1.1–4.0 mm, width 1.0–3.8 mm, length of basal attachment 1.4–3.9 mm. Gall conical, opening round to oblong, 0.2–1.8 mm wide.

(Fig. 18) (n = 15). Body turbinate, margin incised at intersegmental boundaries, length 2.7–3.6 mm, greatest width 1.1–2.3 mm; abdomen tapered, about as long as head + thorax, extending far beyond femur. Eyespots each 15–25 mm wide, on dorsal margin. Antennal segmentation poorly developed; each antenna 80–228 mm long. Frontal lobes difficult to see, each 125–190 µm long, 180–240 µm wide. Tentorial box 300–495 mm long. Pump chamber 34–45 µm long, 33–53 µm wide. Labium 75–160 mm long, 65–220 mm wide. Spiracles 80–125 mm long, 40–75 mm wide across atrium. Fore and mid legs small sclerotic protuberances, 12–63 µm. Hind legs slender and elongate; coxa 270–440 µm long, trochanter + femur 405–630 µm long, tibia 860–1300 µm long, tarsus 250–380 µm long; translucent pores dense throughout dorsal and ventral surfaces of tibiotarsus, a few on distolateral part of coxa; trochanter with 2 campaniform sensilla on each side; femur-tibia articulation non-functional, tibia fixed in orientation parallel to long axis of femur; claw and digitules present but reduced. Anal opening 10–18 µm wide, with anal ring, 18–32 µm wide, sclerotization of ring uneven, weaker at posteroventral end, often appearing horseshoe-shaped, anal ring with 6 minute setae. Anal area with 4 stout spines, 25–35 µm long, each at the end of a fleshy protuberance; one spine on each side of body laterad of anal ring, another on each side of body posterolateral of anal ring, each anterior anal spine with small, auxiliary spine usually present near medial edge of base plus a few minute setae on surface of protuberance.

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.

(Fig. 19) (n = 4). Body outline elliptical, anterior margin incised at midline, length 263–283 µm, greatest width 158–188 µm. Eyespots on margin, each 9–15 mm wide. Antennae 3-segmented, ca 60 mm long, with 4 fleshy setae. Tentorial box 50 mm long. Labium 20–28 mm long, 18–33 mm wide. Spiracles 11–18 mm long, 8 mm wide across atrium. Legs subequal in size: coxa 18–25 µm, with 5 setae, trochanter + femur 53–58 µm long, trochanter with 4 setae, femur with 2 setae, tibia 20–33 µm long, with 4 setae, tarsus 25–35 µm long, with 4 setae, claw 10–13 µm long; tarsal digitules capitate, unequal length, short digitule 18–20 µm long, long digitule 25–28 µm long, claw digitules capitate, each ca 15 µm long. Anal ring 9–15 µm wide, with 6 fine setae, each ca 8 µm long. Apical seta 55–65 µm long.

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.