Austroplebeia magna, Dollin, Anne E., Dollin, Leslie J. & Rasmussen, Claus, 2015

Dollin, Anne E., Dollin, Leslie J. & Rasmussen, Claus, 2015, Australian and New Guinean Stingless Bees of the Genus Austroplebeia Moure (Hymenoptera: Apidae) — a revision, Zootaxa 4047 (1), pp. 1-73 : 60-64

Austroplebeia magna sp. nov.

( Figures 1–11 , 13–25 , 27–29 , 32–33 )

Diagnosis. In workers and males, frons and mesepisternum with plumose hair ( Figs 22 f, 23f, 25g), head width ≥ 1.70 mm ( Figs 10 , 14 ). In workers, pronotal collar and legs without maculations, HBW 0.39–0.47 mm ( Fig. 11 ). Most workers have the following characteristics: thorax markings Grades 0–2 ( Figs 5 , 7 ), plumose hair on clypeus sparser than in A. cassiae , integument moderately visible among hairs ( Fig. 22 f), bristles on basitarsus III brown but bristles on scutellum slightly brown or pale. In males, gonostylus gently curved, thorax markings Grades 5–6 and 10–12, mesepisternum plumose hairs with long barbs at various angles and irregular in shape ( Fig. 25 g), HBW 0.23–0.27 mm ( Fig. 13 ), HTL 1.11–1.19 mm ( Fig. 14 ). In queens, short dense hair covering T6, but absent on T4– T5 ( Fig. 27 b). Distribution NT and far northwestern QLD ( Fig. 28 d).

Description. As in A. cassiae , except for the following characters.

Worker. Holotype: Body 4.0 mm. Forewing (including tegula) 4.0 mm. All other measurements are listed in Table 4 .

Structure. Head width 1.78 mm. Interocellar distance 1.8x ocellorbital distance. Clypeus length 0.5x width. Malar area 0.2x basal mandibular width. Scape length 0.8x alveolus–lateral ocellus distance.

Scutellum length 0.4x width. HBW 0.42 mm.

Colour. Face markings Grade 1. Thorax markings Grade 2, cream.

Pilosity. Density of plumose hair on clypeus moderately sparse. Vertex bristles slightly brown to brown. Long bristles on scutellum slightly brown. Bristles on inner surface of basitarsus III dark brown.

All other workers examined: Body 3.5–4.5 mm. Forewing (including tegula) 4.0– 4.5 mm. All other measurements are listed in Table 4 .

Structure. Head width 1.72–1.90 mm. Interocellar distance 1.5–1.9x ocellorbital distance. Clypeus length 0.5– 0.6x width. Malar area 0.1–0.3x basal mandibular width. Scape length 0.7–0.9x alveolus–lateral ocellus distance.

Scutellum length 0.4x width. Wing venation shown in Fig. 18 . First transverse cubital vein vestige (up to 75 Μm) present in 30% of wings examined and slight trace in 57% (n=53). Vein M of forewing terminates at or before position of anterior end of first recurrent vein in 90% of wings examined (n=54). Cell second Cu of forewing completely indicated by pigmented vein traces in 88% of wings examined (n=40). HBW very broad ( Figs 11 , 19 h), 0.39–0.47 mm.

Sting rudiments. Gonostyli shoulder width moderately broad, 0.12–0.15 mm. Apex of sting stylet usually with a narrowly rounded protrusion. Sting lancet length long, 0.33–0.37 mm ( Figs 15 , 16 h).

Colour. Face markings Grade 1, ranging 0–3a ( Figs 1 , 3 ). Thorax markings Grades 0–2 ( Figs 5 , 7 ), Grade 2 maculations: 56% brown, 44% cream.

Other areas with cream markings: pronotal lobe (sometimes dark brown to yellow-brown); sometimes labrum dull cream. Scape, pronotal collar, metasoma and legs without maculations.

Labrum brown, ranging red-brown to cream. Mandible: orange-brown medially, ranging red-brown to yellowbrown ( Fig. 9 ); dark brown to orange-brown apically. Scape dark brown, sometimes red-brown. Tegula red-brown, ranging dark brown to brown. Basal and medial terga black to red-brown. Apical terga dark brown, ranging to brown ( Fig. 9 ). Sterna usually dark brown.

Pilosity. Density of plumose hair on clypeus much sparser than that on paraocular area—revealing more of surface than in A. cassiae ( Fig. 22 f). Vertex bristles slightly brown or brown.

Long bristles on scutellum slightly brown or pale—finer and paler than in A. cassiae .

In mature, fully coloured specimens: simple hairs of anterior and posterior corbicular fringes of tibia III slightly brown or pale, hairs on apical half of anterior fringe brown or slightly brown, basitarsus III inner surface with coarse brown bristles, deep opaque brown in some regions, becoming slightly brown on eastern end of distribution.

Geographic variation. The extent of thorax colour markings in A. magna sp. nov. workers was 0–4% ( Fig. 10 ), similar to that of A. cassiae . However, some workers near Maningrida in Arnhem Land, NT, had Grade 0 face markings, darker than any observed A. cassiae workers.

In eastern populations, near the southern shore of the Gulf of Carpentaria, thorax colour markings were relatively bright (4%) and vertex, scutellum and basitarsus III bristles were relatively pale.

Male. As in A. cassiae , except for the following characters:

Body 4.0– 4.5 mm. Forewing (including tegula) 4.0– 4.5 mm. All other measurements are listed in Table 5 .

Structure. Head width 1.70–1.82 mm. Interocellar distance 2.4–2.9x ocellorbital distance. Mandible preapical tooth more acutely pointed than A. cassiae but weaker than in A. cincta ( Fig. 17 m). Flagellum relatively longer than in A. cincta , flagellum length 4.1–4.4x scape length.

HBW moderate, 0.23–0.27 mm. HTW moderate, 0.40–0.46 mm ( Fig. 13 ). HTL moderate, 1.11–1.19 mm ( Fig. 14 ).

Colour. Face markings usually Grades 17–18, sometimes 16 ( Figs 2 , 4 ). Thorax markings usually Grades 10– 12, sometimes 5 or 6 ( Figs 5 , 8 ).

Other areas with cream or yellow markings: ventral scape, labrum, all tibiae, all basitarsi (sometimes dull); usually basal mandible, pronotal lobe, a spot on tegula, apical terga and all femora; sometimes pronotal collar (spots on upper edge), metanotum, sterna and trochanters.

Mandible: cream, ranging to red-brown basally; orange-brown to cream medially ( Fig. 9 ). Tegula dark brown to yellow brown, usually with a cream spot.

Pilosity. Mesepisternum plumose hair with barbs longer and sparser than A. cassiae , as in A. australis but varied in angle and overall shape very irregular ( Fig. 25 g).

Queen. Body 5.0– 6.2 mm. Forewing (including tegula) 3.9–4.3 mm.

Structure. HBW 0.25–0.29 mm. Colour. Face markings up to Grade 2 ( Fig. 24 f). Thorax markings up to Grade 4.

Pilosity. Clypeus with moderate density fine hair, mainly simple. Scutellum bristles slightly brown to brown and up to 450 Μm. T1–T5 with sparse fine short hairs, sparse longer (up to 275 Μm) coarse bristles on lateral and dorsal surfaces of T2–T5. T6 with short dense hair ( Fig. 27 b) and an apical fringe of long (up to 375 Μm) simple hairs. Basitarsus III inner surface bristles brown.

Etymology. The Latin feminine adjective, magna , meaning 'large', refers to the broad basitarsus III and long sting lancet in workers of this species.

Material examined. Holotype. ( AMS K.464902, worker), labelled as follows: “ HOLOTYPE / Austroplebeia magna / Dollin, Dollin & / Rasmussen 2015”, “Nest N2 / Worker”, “ AUSTRALIA, NT / 7k SW Katherine / 14.5o S, 132.2o E / 31 July 1987 / Anne & Les Dollin”

Paratypes. ( ANIC 32-069967, worker), labelled as follows: “ PARATYPE / Austroplebeia magna / Dollin, Dollin & / Rasmussen 2015”, “Nest N2 / Worker”, “ AUSTRALIA, NT / 7k SW Katherine / 14.5o S, 132.2o E / 31 July 1987 / Anne & Les Dollin”. ( ANIC 32-069968, male), labelled as follows: “ PARATYPE / Austroplebeia magna / Dollin, Dollin & / Rasmussen 2015”, “Nest N2 / Male”, “ AUSTRALIA, NT / 7k SW Katherine / 14.5o S, 132.2o E / 31 July 1987 / Anne & Les Dollin”. ( AMS K.464903, male), labelled as follows: “ PARATYPE / Austroplebeia magna / Dollin, Dollin & / Rasmussen 2015”, “Nest N2 / Male”, “ AUSTRALIA, NT / 7k SW Katherine / 14.5o S, 132.2o E / 31 July 1987 / Anne & Les Dollin”.

Provenance of types: The holotype and paratypes were all collected from colony N2 of A. magna sp. nov. during an expedition to the NT by Anne and Les Dollin in July–September 1987. Anne and Les Dollin conducted ten major expeditions from 1980 to 2012 to investigate the distribution and characteristics of all species of Australian stingless bees.

Other worker and male specimens examined. Northern Territory: Nest samples, collected by AED & LJD except where otherwise stated: Cobourg Peninsula, 14–19 Aug 1987 (Nest N8, 20 workers, 15 males; Nest N12, 20 workers, 11 males; Nest N14, 10 workers); Buluhkaduru outstation, 50K SE Maningrida, 13–14 Aug 2014 (all by Aung Si: Nest D65, 11 workers, 11 males; Nest D66, 18 workers, 7 males; Nest D67, 28 workers); 7k SW Katherine, 31 Jul 1987 (Nest N1, 20 workers, 2 males; Nest N2, 20 workers, 15 males); Willeroo Station, 120k SW Katherine, Oct 2011 (Nest D57, 20 workers, A. Beil). — Samples collected from flowers or traps: Arnhem Bay, 17 May 1970, N Peterson, AMS, 3 workers; Adelaide R, 28 Sep 1985, AED & LJD, WS-21: 1 worker; 13k W Victoria River, 5 Jan 1989, TF Houston, WAM, 5 workers; Timber Creek, 27 Sep 1985, AED & LJD, WS-19: 4 workers; 68k E Timber Creek, 20 Apr 1980, G Anderson & D Symon, WAM, 2 workers; Roper Bar, 25 Aug 1987, AED & LJD, NS-1: 2 workers; 100k SE Roper Bar, 26 Aug 1987, AED & LJD, NS-3, 2 workers; 260k SE Roper Bar, 28 Aug 1987, AED & LJD, NS-4, 1 worker; Grose Lagoon, 11k SW byS Borroloola, 31 Oct 1975, JC Cardale, ANIC, 12 workers; Caranbirini Waterhole, 33k SW Borroloola, 3 Nov 1975, JC Cardale, ANIC, 6 workers; McArthur River, 14k S byW of Cape Crawford, 25 Oct 1975, JC Cardale, ANIC, 7 workers. Queensland: Sample collected from flowers: 10k NW Westmoreland Station, 20k E of NT/ QLD border, 29 Aug 1987, AED & LJD, NS-7: 15 workers.

Queen specimens examined. Nest samples collected by AED & LJD, 11 queens: Northern territory: Cobourg Peninsula, 14–19 Aug 1987 (Nest N8, 1 virgin; Nest N12, 6 virgin); Katherine, 31 Jul 1987 (Nest N1, 4 virgin).

Remarks. The NT populations here identified as A. magna sp. nov. were part of the 'symei' group defined in our preliminary studies ( Halcroft et al. 2015). That group was characterised by its dark colouration (Thorax Grades predominantly 0–2, Fig. 5 ). It also included the eastern species, A. cassiae .

Although superficially similar to A. cassiae in size and colouration, A. magna workers could be distinguished by their broad basitarsus III, long sting lancet and fine clypeus hair. The structure of the mesepisternum plumose hairs differed in males of A. cassiae and A. magna ( Figs 25 f–g). In addition, the dense layer of hair on T4–T5, seen in queens of A. cassiae , was absent in A. magna ( Fig. 27 ). The broadest basitarsus III observed in the Austroplebeia was in a worker collected from an A. magna colony at Cobourg Peninsula, NT, in the far north of the species range.

Some A. magna workers from near Maningrida, NT, completely lacked pale markings on both the face and thorax. These were the darkest bees observed in the study. All other Austroplebeia workers that we have examined have had at least a dull maculation in the supraclypeal area.

The holotype and paratypes were all collected in 1987 from colony N2 near the middle of the geographic range of this species. The colouration, pilosity characteristics and structural data of the holotype and paratype specimens are all within the ranges described above for A. magna . The holotype (AMS K.464902) and the worker paratype (ANIC 32-069967) are both Face Grade 1, Thorax Grade 2, HBW 0.42 mm ( Figs 1 , 3 , 5 , 7 , 10 , 11 , Table 4 ). One male paratype (AMS K.464903) is Face Grade 17, Thorax Grade 10, HBW 0.24 mm, HTL 1.15 mm; and the other male paratype (ANIC 32-069968) is Face Grade 17, Thorax Grade 5, HBW 0.23 mm, HTL 1.14 mm ( Figs 2 , 4 , 5 , 8 , 13 , 14 ).

*Variables: as given in Table 2.

The following male specimens were measured. A. cincta , N=6 unless otherwise stated, from nests D10 (3x), D14 (3x). A. essingtoni , N=5 unless otherwise stated, from nests N11, N15, W5, W9, W12. A. australis –Central Colour Morph, N=5, from nests C49, N3, Q33, W13, W17. A. australis –Eastern Colour Morph, N=5, from nests C72, C75, F31, NSW55, S2. A. cassiae , N=5, from nests D19, F6, F43, M19, N23. A. magna , N=5, from nests N1, N2, N8, N12 (2x). A. N=5; B. N=4

Figures

FIGURE 1. Face colour marking grades used to classify Austroplebeia workers. In many specimens, these cuticle markings may be obscured by hair on the clypeus and paraocular area. Visualisation of these markings can be improved by tilting the bee's head back and viewing the face obliquely.

FIGURE 2. Face colour marking grades used to classify Austroplebeia males.

FIGURE 3. Variation in worker face colour grades observed within all Austroplebeia taxa except A. cincta. Data show 300 workers from 114 colonies, in addition to 50 workers from 10 flower samples of A. australis – Western Colour Morph. Colour grades are as illustrated in Fig. 1.

FIGURE 4. Variation in male face colour grades observed within all Austroplebeia taxa except A. cincta and A. australis – Western Colour Morph. Data show 186 males from 38 colonies. Colour grades are as illustrated in Fig. 2.

FIGURE 5. Thorax colour marking grades used to classify workers and males of Austroplebeia.

FIGURE 6. Number of different thorax colour marking grades found amongst workers within individual colonies of Austroplebeia. Data are based on 122 colonies and scores of 10 – 49 workers per colony.

FIGURE 7. Variation in worker thorax colour grades observed within all Austroplebeia taxa except A. cincta. Data show 2503 workers from 158 colonies, in addition to 59 workers from 15 flower samples of A. australis – Western Colour Morph. Colour grades are as illustrated in Fig. 5.

FIGURE 8. Variation in male thorax colour grades observed within all Austroplebeia taxa except A. cincta and A. australis – Western Colour Morph. Data show 196 males from 38 colonies. Colour grades are as illustrated in Fig. 5.

FIGURE 9. Variation in colour of the clypeus, mandible (middle part) and apical terga in workers and males of all Austroplebeia taxa. (No males are known in A. cincta NG and A. australis – Western Colour Morph.) Data are based on 120 workers and 62 males.

FIGURE 10. Ratio of head width and extent of thorax colour marking in workers of all Austroplebeia taxa except A. cincta. Data from 121 colonies are shown, in addition to five flower samples from A. australis – Western Colour Morph and one A. australis flower sample with mixed Western / Central characteristics. Data points are averages from 10 – 49 workers per colony, or 5 – 12 workers per flower sample. Data from eight individual type specimens are also shown. The T. symei and T. ornata types, in which the eyes are collapsed preventing accurate head width measurement, are not shown. The T. essingtoni holotype has a missing head, so data from the paratype are shown. 1. Colony F 31 from the northern tip of Cape York Peninsula, QLD, which has characteristics similar to A. australis – Eastern Colour Morph despite its geographic isolation. Names given in brackets in the legend are groups in our Halcroft et al. (2015) preliminary analysis.

FIGURE 11. Ratio of tibia III width and basitarsus III width in workers of all Austroplebeia taxa. Data from 173 colonies are shown, in addition to 26 flower samples: A. australis – Western Colour Morph (14 x), A. australis with mixed Western / Central characteristics (4 x) and NG A. cincta (8 x). Data points are averages from 2 – 4 workers per colony, or 1 – 12 workers per flower sample. Data from 12 individual type specimens are also shown. 1. Colony F 31 from the northern tip of Cape York Peninsula, QLD, which has characteristics similar to A. australis – Eastern Colour Morph despite its geographic isolation. Names given in brackets in the legend are groups in our Halcroft et al. (2015) preliminary analysis.

FIGURE 13. Ratio of tibia III width and basitarsus III width in males of all Austroplebeia species. Data points are averages from 2 – 3 males per colony. Data from the male paratypes of A. magna sp. nov. are also shown. Names given in brackets in the legend are groups in our Halcroft et al. (2015) preliminary analysis.

FIGURE 14. Ratio of head width and tibia III length in males of all Austroplebeia species. Data points are averages from 2 – 3 males per colony. Data from the male paratypes of A. magna sp. nov. are also shown. Names given in brackets in the legend are groups in our Halcroft et al. (2015) preliminary analysis.

FIGURE 15. Worker sting rudiments metric data: ratio of maximum length of sting lancet (A) and shoulder width of gonostyli (B) in all Austroplebeia taxa. Data points are averages for 1 – 3 workers per nest or flower sample. 1. The geographically isolated colony, F 31, from the northern tip of Cape York Peninsula, QLD, which has colouring and HTW / HBW similar to A. australis – Eastern Colour Morph. 2. Colonies C 58 and D 55 from the ‘ intermediate’ group of Halcroft et al. (2015) which are consistent with A. australis – Eastern Colour Morph in this analysis. 3. CS- 8 workers, with Grade 4 Thorax Colour Markings, collected from flowers in the MacDonnell Ranges, NT. They are consistent with A. australis – Central Colour Morph in this analysis, despite their relatively dark colouration. 4. GS- 10 workers collected from flowers on the far western edge of the A. cassiae range. 5. NS- 7 workers collected from flowers on the far eastern edge of the A. magna sp. nov. range. Names given in brackets in the legend are groups in our Halcroft et al. (2015) preliminary analysis.

FIGURE 16. Austroplebeia worker sting rudiments: a, A. cincta NG; b. A. cincta QLD; c, A. essingtoni; d, A. australis – Eastern Colour Morph; e, A. australis – Central Colour Morph; f, A. australis – Western Colour Morph; g, A. cassiae; h, A. magna sp. nov. All drawn to same scale. Abbreviations: Gon—gonostylus; Sty—sting stylet (bulb); Lan—sting lancet (first valvula or first ramus).

FIGURE 17. Austroplebeia antennae, mandibles and labial palpi. Figs a – c. Antenna (basal part) of A. essingtoni: a, worker; b, male; c, queen. Figs d – h. Left mandible of worker: d, A. cincta QLD; e, A. essingtoni; f, A. australis; g, A. cassiae; h, A. magna sp. nov. Figs i – m. Left mandible of male: i, A. cincta QLD; j, A. essingtoni; k, A. australis; l, A. cassiae; m, A. magna sp. nov. Fig. n. Left mandible of A. australis queen. Figs a – n drawn to same scale. Figs o – q. Labial palpus of worker (mesal margins uppermost, small hairs omitted): o, A. cincta; p, A. cassiae; q, A. magna sp. nov. Figs o – q drawn to same scale. Abbreviations: F 1 — first flagellomere; F 2 — second flagellomere; F 3 — third flagellomere; F 2 L—second flagellomere length (showing how this measurement was made).

FIGURE 18. Austroplebeia wings. Dotted lines indicate pigmented vein traces. Figs a – c. Forewings, showing variation observed in the vestige of the first transverse cubital vein: a, A. magna sp. nov. forewing with a short vein vestige; b, Detail of an A. australis – Eastern Colour Morph forewing with a thickening of vein M at this position; c, Detail of an A. australis – Eastern Colour Morph forewing with a long vein vestige. Fig. d. Hindwing of A. magna sp. nov. Fig. e. Diagram of a forewing showing how the following measurements were made: (1) forewing length without tegula; (2) forewing width; (3) wing diagonal (Sakagami 1978); (4) 1 st abscissa of M length; (5) 1 st abscissa of Cu length. Figs a – d drawn to same scale: scale bar = 0.25 mm. Abbreviations: M—vein M; 1 st R—first recurrent vein; 2 nd Cu—second cubital cell; Ham—hamuli.

FIGURE 19. Austroplebeia legs. Figs a – b. Worker tibia III and basitarsus III of A. australis – Central Colour Morph: a, inner surface; b, outer surface. Fig. c. Cross-sectional view of A. australis worker tibia III, inner surface uppermost. Fig. d. Worker leg II of A. australis – Eastern Colour Morph, showing mesotibial spur near junction of tibia and basitarsus. Figs e – f. Male tibia III and basitarsus III of A. australis – Central Colour Morph: e, inner surface; f, outer surface. Fig. g. Queen tibia III and basitarsus III of A. australis – Central Colour Morph, outer surface. Fig. h. Variation in size of worker basitarsus III (shaded segment), all drawn to same scale: A. cincta QLD is shown on the left, then remaining species are shown in order of increasing basitarsus III width. Abbreviations: Kei—keirotrichia; Pos—posterior corbicular fringe; Ant—anterior corbicular fringe; Rasrastellum; Cor—corbicula; Pen—pencillum.

FIGURE 20. Austroplebeia male sterna and terga, shown in ventral view except for Fig. r. Figs a – f. S 4: a, A. cincta; b, A. essingtoni; c, A. australis; d – e, A. cassiae showing variation in position of gradulus; f, A. magna sp. nov. Figs g – j. S 5: g, A. cincta; h, A. essingtoni, arrow indicates protruding apical lobe; i, A. australis; j, A. cassiae. Figs k – o. S 6: k, A. cincta; l, A. essingtoni; m, A. australis; n, A. cassiae; o, A. magna sp. nov. Figs p – q. S 7: p, A. cincta; q, A. australis. Fig. r. T 7 of A. cassiae, dorsal view. Figs a – f, scale bar = 0.5 mm; figs g – r, scale bar = 0.33 mm. Abbreviations: Gra—gradulus; Med—median body.

FIGURE 21. Austroplebeia male genitalia. Figs a – c. Genitalia, dorsal view on left, ventral view on right: a, A. cincta; b, A. essingtoni; c, A. magna sp. nov. Figs d – e. Apical abdomen and protruding parts of genitalia, viewed laterally, as visible in many pinned male specimens: d, A. essingtoni, showing distinct curve in apical gonostylus; e. A. magna sp. nov., with gently curved apical gonostylus. Figs a – c drawn to same scale. Abbreviations: Pen—penis valve; Gon—gonostylus.

FIGURE 22. Austroplebeia worker faces: a, A. cincta NG; b, A. cincta QLD; c, A. essingtoni, showing a relatively heavily marked specimen within the species range; d, A. australis – Eastern Colour Morph; e, A. cassiae; f, A. magna sp. nov. Scale bars = 0.25 mm.

FIGURE 23. Austroplebeia male faces: a, A. cincta QLD; b, A. essingtoni; c, A. australis – Eastern Colour Morph; d, A. australis – Central Colour Morph; e, A. cassiae; f, A. magna sp. nov. Scale bars = 0.25 mm.

FIGURE 24. Austroplebeia queen faces: a, A. cincta QLD; b, A. essingtoni; c, A. australis – Eastern Colour Morph; d, A. australis – Central Colour Morph; e, A. cassiae; f, A. magna sp. nov. Scale bars = 0.25 mm.

FIGURE 25. Austroplebeia lateral thoraces: Figs a – b. Worker mesosomal side: a, A. cincta QLD; b, A. cassiae. Figs c – d. Male mesosomal side: c, A. cincta QLD; d, A. cassiae. Figs e – g. Detail of mesepisternum pilosity on male mesosomal side (photographs by K. Walker): e, A. australis; f, A. cassiae; g, A. magna sp. nov. Figs e – g shown at same magnification; all scale bars = 0.25 mm. Abbreviation: Met—metepisternum.

FIGURE 27. Pilosity on apical part of Austroplebeia queen abdomen, viewed laterally: a, A. cassiae; b, A. magna sp. nov. Scale bars = 0.25 mm.

FIGURE 28. Observed distribution of all Austroplebeia species in Australia and the island of New Guinea: a, A. cincta; b, A. essingtoni; c, A. australis; d, A. magna sp. nov. and A. cassiae. In all maps, square symbols represent nest samples and circles represent samples collected from flowers or traps. The type locality for each named species, including those now synonymised, is indicated with arrows. Where the type locality was a general area such as ' Central Australia' or ' Cape York', these areas are indicated with shading.

FIGURE 29. Austroplebeia nest entrances: a, A long A. cincta tunnel (120 mm) near Rossville, QLD; b, The tunnel of an A. essingtoni colony that has nested in a cliff face crevice 65 k S of Maningrida, NT (photograph by Aung Si); c, A translucent fresh tunnel of an A. australis colony near Tara, QLD (photograph by M. Halcroft); d, An A. australis colony without an entrance tunnel, near Tara, QLD (photograph by M. Halcroft); e, A thicker and older tunnel of an A. cassiae colony near Rossville, QLD; f, The tunnel of an A. magna colony 50 k S of Maningrida, NT (photograph by Aung Si); g, The longest tunnel recorded in this study: the 430 mm tunnel of an A. cincta colony that was being harassed by green ants near Rossville, QLD; h, Another long tunnel (ca. 120 – 130 mm) of an A. magna sp. nov. colony 50 k S of Maningrida, NT. Green ants can also be seen on this tunnel (photograph by Aung Si); i, The 160 mm tunnel of an A. cassiae colony near Rossville, QLD. The green ants seen on this tunnel have caught two forager bees.

FIGURE 32. Internal nest characteristics and brood combs of species other than A. cincta: a, Nest of an A. essingtoni colony at Broome, WA (photograph by T. Heard); b, Brood comb of an A. australis colony in a natural log near Ti Tree, NT; c, Brood comb of a boxed colony of A. cassiae near Rossville, QLD; d, Brood comb of an A. magna sp. nov. colony in a narrow tree cavity, 50 k S of Maningrida, NT (photograph by Aung Si). Abbreviations: Lar—larval brood cells; Coc—cocoons; Pol—pollen pots; Hon—honey pots; Bat—batumen layer.

FIGURE 33. Honey and pollen storage pots in Austroplebeia colonies: a, Translucent honey pots in an A. cincta colony near Rossville, QLD; b, Honey pots with thick, strongly pitted walls in an A. australis colony near Tara, QLD (photograph by M. Halcroft); c, Honey pots in an A. magna sp. nov. colony 50 k S of Maningrida, NT (photograph by Aung Si); d, Pollen pots in an A. cincta colony near Rossville, QLD; e, Pollen pots with thickened pitted walls in an A. australis colony near Tara, QLD (photograph by M. Halcroft); f, Pollen pots with strongly pitted walls in an A. australis colony near Kilcoy, QLD; g, Pollen pots made from red-brown cerumen in an A. cassiae colony at Conjuboy, QLD.

Tables

TABLE 5. Structural dimensions from Austroplebeia males. Measurements in mm.

VAR*	A. cincta QLD Mean SD	A. essingtoni Mean SD	A. australis Colour Morphs Eastern Central Mean SD Mean	SD	A. cassiae Mean SD	A. magna Mean SD
1 2 3	3.88 0.17 1.56 0.02 1.27 0.02	3.82 0.30 1.57 0.03 1.25 0.04	4.43 0.38 4.24 1.81 0.04 1.69 1.43 0.04 1.35	0.18 0.04 0.04	4.31 0.17 1.79 0.05 1.43 0.04	4.26 0.24 1.77 0.04 1.39 0.03
4 5 6	1.09 0.02 0.87 0.01 0.92 0.01	1.06 0.01 0.88 0.03 0.91 0.03	1.21 0.03 1.12 1.04 0.03 0.96 1.07 0.03 0.99	0.03 0.04 0.03	1.22 0.02 1.03 0.03 1.07 0.04	1.19 0.03 1.03 0.02 1.06 0.02
7 8 9	0.61 0.01 0.17 0.01 0.35 0.01	0.61 0.02 0.18 0.00 0.39 0.01	0.71 0.02 0.69 0.19 0.01 0.18 0.44 0.01 0.44	0.03 0.01 0.02	0.74 0.01 0.20 0.01 0.42 0.01	0.74 0.01 0.20 0.01 0.43 0.01
10 11 12	0.13 0.01 0.18 0.02 0.65 0.02	0.12 0.01 0.17 0.02 0.61 0.02	0.18 0.02 0.15 0.20 0.03 0.19 0.66 0.02 0.61	0.01 0.02 0.03	0.17 0.01 0.19 0.03 0.67 0.03	0.16 0.01 0.19 0.02 0.65 0.04
13 14 15	0.15 0.01 0.35 0.02 0.58 0.01	0.16 0.02 0.34 0.02 0.56 0.02	0.17 0.03 0.16 0.41 0.04 0.39 0.66 0.02 0.62	0.02 0.01 0.02	0.18 0.02 0.39 0.02 0.68 0.02	0.18 0.01 0.39 0.02 0.70 0.03
16 18 19	0.79 0.01 0.20 0.01 1.54 0.04	0.77 0.02 0.20 0.01 1.84 0.07	0.87 0.01 0.81 0.23 0.01 0.22 2.06 0.06 1.95	0.02 0.01 0.04	0.88 0.01 0.25 0.01 2.06 0.05	0.83 0.03 0.21 0.01 2.01 0.02
20 21 22	0.41 0.01 0.13 0.00 0.11 0.01	0.43 0.01 0.13 0.01 0.11 0.01	0.48 0.02 0.45 0.14 0.01 0.14 0.12 0.02 0.12	0.01 0.01 0.01	0.49 0.02 0.14 0.01 0.13 0.01	0.47 0.02 0.14 0.01 0.12 0.01
23 24 25	0.08 0.01 0.14 0.01 0.16 0.01	0.08 0.01 0.15 0.01 0.18 0.02	0.09 0.01 0.09 0.18 0.01 0.16 0.21 0.02 0.19	0.01 0.01 0.01	0.09 0.02 0.16 0.01 0.19 0.01	0.09 0.01 0.16 0.01 0.21 0.01
26 27 28 29 30	0.13 0.01 0.14 0.02 0.12 0.01 1.04 A 0.03 1.15 0.03	0.13 0.01 0.17 0.01 0.14 0.01 1.05 0.02 1.13 0.02	0.16 0.01 0.15 0.20 0.01 0.18 0.16 0.01 0.15 1.23 0.04 1.17 1.31B 0.02 1.25	0.01 0.01 0.01 0.05 0.03	0.16 0.01 0.19 0.01 0.16 0.01 1.23 0.04 1.37 0.05	0.15 0.00 0.18 0.01 0.15 0.00 1.22 0.05 1.35 0.01
31 32 33 34 35 36 37 38	0.30 0.03 0.64A 0.04 3.17 0.07 3.59 0.09 1.25 0.04 0.94 0.03 0.49 0.02 0.70 0.04	0.32 0.00 0.72 0.02 3.18B 0.04 3.61B 0.08 1.29B 0.03 0.90B 0.02 0.45B 0.03 0.69B 0.03	0.34 0.02 0.35 0.80 0.03 0.81 3.77B 0.16 3.55 4.35B 0.16 4.06 1.49B 0.07 1.44 1.06 0.06 1.04 0.55 0.02 0.52 0.82 0.02 0.80	0.03 0.03 0.11 0.16 0.05 0.05 0.04 0.07	0.35 0.02 0.84 0.05 3.70 0.14 4.34 0.17 1.45 0.04 1.06 0.06 0.55 0.05 0.82 0.04	0.37 0.01 0.87 0.02 3.67 0.07 4.24 0.17 1.51 0.05 1.09 0.03 0.55 0.02 0.79 0.03
39 40	5.00 0.00 1.09 0.02	5.00 0.00 1.04 0.02	5.80 0.45 5.80 1.27 0.06 1.19	0.45 0.02	5.40 0.55 1.26 0.04	5.40 0.55 1.17 0.01
41 42	0.37 0.02 0.20 0.01	0.39 0.01 0.21 0.01	0.49 0.03 0.43 0.27 0.01 0.23	0.01 0.02	0.46 0.02 0.26 0.01	0.43 0.02 0.24 0.01