Austroplebeia australis

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 : 46-53

Austroplebeia australis –Western Colour Morph

Diagnosis. As in general diagnosis of A. australis with the following exceptions:

Worker HBW 0.33–0.40 mm ( Fig. 11 ). Most workers have the following characteristics: thorax markings Grades 4, 9 or 10 ( Figs 5 , 7 ), ventral scape dark brown or red-brown, often with a cream maculation, upper edge of pronotal collar maculations limited to two small lateral spots, apical terga cream ( Fig. 9 ), leg maculations limited to a basal spot on tibia I. Male and queen unknown. Distribution northern areas of The Kimberley, WA ( Fig. 28 c).

Description. As in A. australis –Eastern Colour Morph, except for the following characters.

Worker. Body 3.7–4.7 mm. Forewing (including tegula) 4.1–4.5 mm. All other measurements are listed in Table 3 .

Structure. Head width 1.72–1.88 mm. Interocellar distance 2.0–2.3x ocellorbital distance. Scape length 0.8– 0.9x alveolus–lateral ocellus distance.

Wing venation similar to that shown in Fig 18 . First transverse cubital vein vestige (up to 90 Μm) present in 38% of wings examined and slight trace in 54% (n=48). Vein M of forewing terminates at or before position of anterior end of first recurrent vein in 77% of wings examined (n=47). Cell second Cu of forewing completely indicated by pigmented vein traces (n=44). Hamuli 6 per wing. HBW moderately broad ( Fig. 11 ), 0.33–0.40 mm.

Sting rudiments. Gonostyli shoulder width broad, 0.14–0.17 mm. Apex of sting stylet angular or broadly rounded. Sting lancet length moderately long, 0.29–0.32 mm ( Figs 15 , 16 f).

Colour. Face markings usually Grade 3, ranging 1–4 ( Figs 1 , 3 ). Thorax markings usually Grades 4, 9 or 10, ranging 2–11 ( Figs 5 , 7 ).

Other areas with cream or yellow markings: pronotal lobe, upper edge of pronotal collar (two to four spots); usually apical terga, a spot on tegula, and a basal spot on tibia I; sometimes labrum and ventral scape.

Labrum usually brown, sometimes dull cream. Scape dark brown to red-brown, with varying amounts of cream ventrally. Flagellum ranging orange-brown to yellow-brown. Tegula red-brown or brown, usually with a cream spot. Basal and medial terga usually black to red-brown. Apical terga usually cream, sometimes red-brown to yellow-brown ( Fig. 9 ). Unmarked areas of tibiae dark brown to orange-brown. Trochanter red-brown or orangebrown. Mediotarsus usually dark brown.

Pilosity. As in A. australis –Eastern Colour Morph.

Geographic variation. The extent of thorax colour markings in the Western Colour Morph was generally 9– 13% ( Fig. 10 ). Scores of HTW/HBW were generally larger in the more northerly populations of their range.

Worker bees with mixed characteristics were found between the ranges of the Western and Central Colour Morphs ( Fig. 28 c), with more extensive thorax markings (17%) but with HTW/HBW scores within the Western Colour Morph range.

Male and queen. Unknown.

Material examined. Lectotype of Trigona australis . ( ZMHB, worker), labelled as follows: “C. Austr. / v. Müller.93.”, “ Trigona [worker symbol] / australis / det. Friese 1898 / n.sp.”, “ LECTOTYPE / Trigona / australis Friese / Designated C Rasmussen’07”.

Provenance of types: The lectotype specimen is labelled as originating from Central Australia where it presumably was collected by Ferdinand von Müller (1825–1896) in 1893. However, it is likely that the type specimen was actually collected by the Hermannsburg missionary, Friedrich Adolf Hermann Kempe (1844–1928), who sent insect specimens to von Müller in Melbourne (John Strehlow, pers. comm. to AED, 1996). Specimens collected by Kempe came from the area around Hermannsburg and Finke River in the NT where Kempe resided from 1877 to December 1891. The lectotype specimen would therefore have been collected no later than 1891 and perhaps 1893 was the time when Müller received and labelled the specimens with his name. Additional evidence that the collector would not have been von Müller is from Chris Torlach of the Strehlow Research Centre in Alice Springs, NT (pers. comm. to AED, 1996). She said she had no reports of visits to the NT by von Müller, but that von Müller had contacts there making collections for him. The lectotype selected was from Friese’s own collection now in ZMHB.

Additional paralectotype material includes specimens from Burnett River, QLD, collected by Richard Wolfgang Semon (1859–1918) in 1891–92 (expedition details in Semon (1894)). Friese mentioned numerous specimens (quantity unknown) from von Müller and several specimens from Semon in the collections of Stuttgart, Amsterdam and Budapest. A paralectotype was seen from Friese’s own collection now in ZMHB. The specimen was not examined in detail but its labels are as follows: “Burnett River / Queensland / 1891/92. R. Semon”, “ Trigona [worker symbol] / australis / det. Friese 1897 / n.sp.”, “ PARALECTOTYPE / Trigona / australis Friese / Designated C Rasmussen’07”.

Lectotype of Trigona percincta . Here designated ( ZMHB, worker), labelled as follows: “I. Australien / Hermannsburg / Finke River / v. Leonhardi lg., “ Trigona cincta Mocs. var. [identification label in T.D.A. Cockerell’s handwriting], “ LECTOTYPE / Trigona percincta Cockerell , / Design. C Rasmussen 2013.

Provenance of types: A series of four specimens, labelled as from Moritz Freiherr von Leonhardi (1856–1910) and collected at Hermannsburg, Finke River, NT, had been identified in ZMHB as a possible variety of Trigona cincta by Friese (unpublished) and Cockerell (1910). Later, Cockerell (1929a) corrected this previous identification of 'four specimens from Australia (Hermannsburg, Finke River, V. Leonhardi)' and named the species Trigona cincta percincta . Cockerell, however, did not revise the specimens again and they were never labelled as percincta . The four specimens are now labelled as lectotype and paralectotypes, respectively, and based on the information from Cockerell, they are unmistakably the material on which Cockerell based his description. One of the paralectotypes is in addition labelled ' Trigona / cincta ? / [worker symbol] 1909 Friese det. / juven. Mocs.' (name and last line in Friese’s handwriting).

Although the specimen came to ZMHB from von Leonhardi, it is likely that the actual collector was the Hermannsburg missionary Carl Friedrich Theodor Strehlow (1871–1922). John Strehlow (pers. comm. to AED, 1996) explained that Carl Strehlow received a letter from von Leonhardi (dated 23 Oct 1908) requesting 'honeygathering bee species in the area'. The request from von Leonhardi was repeated in the next letter dated 23 Dec 1908. It is known that insects were collected by the Aranda Aboriginals living near the mission, who got paid for it. They probably located a stingless bee nest and collected several specimens for Strehlow. From further letters in John Strehlow's collection, it is clear that bee samples were sent from Hermannsburg in 1909 and von Leonhardi identified them first as ' Trigona australis ' (letter dated 31 Oct 1909). The specimens were then probably forwarded to Senckenberg Museum in Frankfurt, Germany, for positive identification. In the next letter from von Leonhardi (dated 14 Nov 1909), the name was updated to [what appears to be] ' Trigona cincta '. In the letter it is described how the colour was not fully developed due to the young age of the specimens, which Friese also indicated on his identification label (as juven. for juvenile or callow). Presumably the specimens were transferred from Frankfurt to ZMHB, where they are now.

Holotype of Trigona cockerelli . ( ANIC, worker), labelled as follows: “ TYPE, “Borroloola, N.T. / 25.8.1911 / G. F. Hill, “ Trigona / cockerelli Raym / [reverse] Euryglossa strigata Ckll, “ AUST. NAT. / INS. COLL.

Paratype of Trigona cockerelli . ( ANIC worker), labelled as follows: “Borroloola, N.T. / 25.8.1911 / G. F. Hill, “ PARATYPE, “ ANIC

Provenance of types: The holotype was collected by Gerald Freer Hill (1880–1954) on 25 August 1911 (listed as September by Rayment 1930) in Borroloola, NT. Hill was a naturalist for the Commonwealth Government Exploration Party, Central Australia and Northern Territory from 1911 to 1912 and his insect collection was deposited in ANIC where Rayment located and described Trigona cockerelli . An additional paratype is found in ANIC. Another worker of same collection event is in NMV, listed in 'Other specimens examined' for the Central Colour Morph, below.

Lectotype of Trigona cockerelli ornata . ( NMV, worker), designated Rasmussen 2008, labelled as follows: 4, “ TYPE, “ Trigona / cockerelli RAYM . / ORNATA / C. YORK Q., “ SYNTYPE / T-11575 / Trigona / cockerelli / ornata , “MUS. VIC. / ENTO 2008-8 L

Paralectotypes of Trigona cockerelli ornata . ( NMV, worker), designated Rasmussen 2008, labelled as follows: 4, "HYM-34695, “ PARALECTOTYPE / T-21946 / Trigona cockerelli ornata , “MUS. VIC. / ENTO 2014-18 L; ( NMV, worker), designated Rasmussen 2008, labelled as follows: 4, "HYM-34696, “ PARALECTOTYPE / T- 21947 / Trigona cockerelli ornata , “MUS. VIC. / ENTO 2014-18 L; ( NMV, worker), designated Rasmussen 2008, labelled as follows: 4, "HYM-34697, “ PARALECTOTYPE / T-21948 / Trigona cockerelli / ornata , “MUS. VIC. / ENTO 2014-18 L; ( ANIC, worker), labelled as follows: “Cotype, “ Trigona ornata / cockerelli RAYM / Cape York. Q. / DONALD THOMPSON, “ AUST. NAT. / INS. COLL; ( ANIC, worker), labelled as follows: “Cotype, “ AUST. NAT. / INS. COLL; ( ANIC, male), labelled as follows: “Allotype, “ Trigona [male symbol] RAYM / cockerelli ornata / C. York. Q., “ AUST. NAT. / INS. COLL;

Provenance of types: According to the type description ( Rayment 1935), the T. cockerelli ornata specimens were collected by Donald Thomson, D.Sc. on Cape York. This is a peninsula covering nearly 140,000 km 2 in far north QLD. However, it is possible to narrow down the collection locality of these specimens. Prior to the first publication of this species name in Rayment (1932), Donald Finlay Fergusson Thomson (1901–1970) had visited less than 30% of Cape York in his field work. Thomson had conducted two overland expeditions in 1928 and 1929 in central Cape York, between Iron Range (12.8o S x 143.3o E) in the northeast and Kowanyama (15.5o S x 141.75o E) in the southwest ( Thomson 1935). These specimens were probably collected in this area.

The lectotype and three paralectotype specimens are in NMV, but two additional paralectotype (as 'cotype') workers and one (as 'allotype') male are located in ANIC. According to Rayment (1935) the specimens were sent directly from Thomson to Rayment for identification.

Holotype of Trigona websteri . ( ANIC, worker), labelled as follows: “ TYPE, “ Trigona / websteri Raym. / WYNDHAM, W.A. / [reverse] 25/1/1931, “ AUST. NAT. / INS. COLL.

Paratype of Trigona websteri . ( ANIC, worker), labelled as follows: “Cotype, “ Trigona / websteri Raym. / WYNDHAM, W.A. / [reverse] 25/1/1931, “ ANIC.

Provenance of types: The holotype was collected by Victor Henry Webster (1905–1981), a medical doctor, on 25 Jan 1931 in Wyndham, WA. According to his son, John Webster (pers. comm. to AED, 2011), Webster senior had a good relationship with the local Aboriginal people and could have collected the bee specimens together with them. John knew Webster would sometimes go with others on trips to the surrounding countryside in the rickety two wheel drives of the time. According to Rayment (1935) the specimens were sent directly from Webster to Rayment for identification. The date of the collection could be earlier, as the medical staff and others returned to Perth each year during the wet season (including January). Either Victor Webster remained in Wyndham that year during the wet season or the date was the date of labelling. The holotype and a single paratype are deposited in ANIC.

Other worker and male specimens examined.

Austroplebeia australis –Eastern Colour Morph. Queensland: Nest samples, collected by AED & LJD except where otherwise stated: Cape York near Bamaga, 10 Aug 1983 (Nest F31, 20 workers, 4 males); Princhester, 5 Oct 1996 (Nest C75, 20 workers, 2 males); Rockhampton, 8–9 Sep 1996 (Nest C26, 14 workers; Nest C27, 20 workers; Nest C30, 13 workers); Duaringa, 31 Oct 1987 (Nest N27, 5 workers, E. Adams); Sep 1996 (Nest C32, 20 workers; Nest C33, 19 workers; Nest C34, 17 workers; Nest C35, 20 workers; Nest C39, 20 workers); Oct 1996 (Nest C56, 18 workers; Nest C63, 5 workers; Nest C67, 20 workers; Nest C70, 20 workers; Nest C72, 5 workers, 20 males; Gin Gin, 5 Sep 1996 (Nest C11, 3 workers), 19 Oct 1996 (Nest C83, 9 workers); Tin Can Bay, 3 Sep 1996 (Nest C3, 5 workers); from Rainbow Beach but kept at Ipswich, Sep 2013 (Nest D35, 31 workers, C. Heather); Kilcoy, Oct 1989 (Nest S2, 20 workers, 5 males; Nest S3, 17 workers; Nest S5, 8 workers; Nest S6, 20 workers; Nest S9, 12 workers; Nest S34, 16 workers); Kenilworth, Oct 2013 (Nest D48, 10 workers, D Clarke); Blackbutt, 24 Oct 1989 (Nest S29, 5 workers; Nest S31, 15 workers); Miles, 2 Sep 2013 (Nest D37, 9 workers, R. Raymond; Nest D38, 10 workers, R. Raymond); Tara, Sep 2013 (Nest D39, 15 workers; R. Raymond; Nest D40, 28 workers, 3 males, A. Beil; Nest D41, 49 workers, A. Beil; Nest D42, 39 workers, A. Beil;); Gold Coast, Pimpama, Jan 2014 (Nest D53, 20 workers, P Davenport); Gold Coast, Guanaba, Jan 2014 (Nest D56, 20 workers, P Davenport); Gold Coast, Bonogin, Jan 2014 (Nest D55, 20 workers, P Davenport); Gold Coast, Tallebudgera, Jan 2014 (Nest D54, 20 workers, P Davenport); Herries Range near Karara, 26 Oct 1989 (Nest S33, 14 workers, 3 males); Eulo, 4 Mar 1982 (Nest Q35, 20 workers). — Samples collected from flowers or traps: Dalby, 8 Jan 1984, K Walker, NMV, 4 workers; 40k S Dalby, 10 Jan 1984, K Walker, NMV, 2 workers; 5k N Karara, 6 Jan 1984, K Walker, NMV,> 20 workers; 10k S Leyburn, 6 Jan 1984, K Walker, NMV, 6 workers. New South Wales: Nest samples, collected by AED & LJD except where otherwise stated: Lismore, 23 Jan 1980 (Nest 33, 10 workers); Ashford, Dec 2013 (Nest D51, 20 workers, A Beil); Kempsey, Jan 1980 (Nest 15, 53 workers; Nest 18, 2 males; Nest 21, 20 workers, 2 males; Nest 23, 40 workers; Nest 24, 10 workers; Nest 25, 10 workers, 3 males; Nest 42, 2 workers, 12 males); Feb 1980 (Nest 50, 2 workers; Nest 52, 5 workers; Nest 53, 2 workers; Nest 54, 20 workers; Nest 55, 10 workers, 4 males; Nest 56, 5 workers); 26 Oct 1980 (Nest 67, 20 workers;); Loomberah near Tamworth, 14 Jan 1980 (Nest 10, 2 workers); Liverpool Ranges near Merriwa, 16 Dec 1979 (Nest 1, 3 workers); Dungog, 18 Jan 1980 (Nests 80, 83, 85, 87, 88, 89, 90, 92 and 93, workers examined and nests photographed by AED & LJD, but specimens not collected). — Samples collected from flowers or traps: Mill-Bullah Waterholes, Waa Gorge, Kaputah NP, 44.2k NE Narrabri, 13 Nov 2009, DR Britton & J Recsei, AMS, 8 workers; Wootton, 24 Feb 2002, M Batley, AMS, 1 worker.

Austroplebeia australis –Central Colour Morph. Queensland: Nest samples, collected by AED & LJD except where otherwise stated: Koolatah Station, 23–24 Sep 1998 (Nest M17, 20 workers; Nest M22, 14 workers); 15k NW of Dunbar Station, 24 Aug 1990 (Nest G8, 15 workers); Delta Downs Station, Sep 2013 (Nest D44, 22 workers, A. Beil); Croydon, 31 Aug 1987 (Nest N21, 20 workers), 19–20 Aug 1990 (Nest G1, 20 workers; Nest G2, 6 workers); Hughenden, 28 Feb 1982 (Nest Q33, 5 workers, 3 males; Nest Q34, 5 workers), 30–31 Aug 1983 (Nest F52, 20 workers; Nest F53, 20 workers). — Samples collected from flowers or traps: M Lagoon, 35k E Wernadinga Station, 30 Aug 1987, AED & LJD, NS-9: 5 workers; 70k N Mt Isa, 18 Sep 1996, AED & LJD, CS-6: 4 workers; 20k E Mt Isa, 6 May 2008, M Batley, AMS, 1 worker; Cloncurry, 17 Oct 1953, NMV, 3 workers; 16k W Cloncurry, 6 Aug 2012, M Batley, AMS, 3 workers; 5k SW Cloncurry, 17 Sep 1996, AED & LJD, CS-5: 4 workers; Hughenden, 30 Aug 1983, AED & LJD, 3 males; 140k N Clermont, 16 Apr 2005, M Batley, AMS, 1 worker, 108k W Winton, 19 Apr 2013, M Batley, AMS, 2 workers. Northern Territory: Nest samples, all collected by AED & LJD except where otherwise stated: Katherine, 31 Jul 1987 (Nest N3, 20 workers, 15 males); 50k W Timber Creek, 27 Sep 1985 (Nest W13, 14 workers, 7 males; Nest W14, 10 workers; Nest W15, 4 workers; Nest W16, 8 workers; Nest W17, 20 workers, 11 males); Borroloola, 28 Aug 1987 (Nest N19, 3 workers); Utopia, Jun 2015 (Nest D73, 10 workers, 3 males, Aung Si & M Turpin; Nest D74, 17 workers, Aung Si & M Turpin); Ti Tree, Jun 2015 (Nest D72, 12 workers, 3 males, Aung Si & M Turpin); 60k S Ti Tree, 3 Oct 1996 (Nest C47, 20 workers; Nest C48, 20 workers; Nest C49, 20 workers, 6 males; Nest C50, 5 workers; Nest C51, 20 workers; Nest C52, 5 workers; Nest C53, 20 workers). — Samples collected from flowers or traps: 13k W Victoria River, 5 Jan 1989, TF Houston, WAM, 5 workers; 42k W Timber Creek, 4 Apr 1980, G Anderson & D Symon, WAM, 1 worker; 35k W Timber Creek, 20 Apr 1980, G Anderson & D Symon, WAM, 1 worker; 56k E Timber Creek, 20 Apr 1980, G Anderson & D Symon, WAM, 1 worker; 68k E Timber Creek, 20 Apr 1980, G Anderson & D Symon, WAM, 1 worker; 50k WSW Wave Hill 29 Apr 1973, CA&TF Houston, WAM, 1 male; Borroloola, 25 Aug 1911, GF Hill, NMV, 1 worker; Grose Lagoon, 11k SW byS Borroloola, 31 Oct 1975, JC Cardale, ANIC, 1 worker; Caranbirini Waterhole, 33k SW Borroloola, 3 Nov 1975, JC Cardale, ANIC, 15 workers; McArthur River, 14k S byW of Cape Crawford, 25 Oct 1975, JC Cardale, ANIC, 1 worker; 83k E Top Springs, 26 Jan 2004, S Cowan, AMS, 1 worker; Elliot, 13 May 2008, M Batley, AMS, 1 worker; 130k N Tennant Creek, 1 Apr 2008, G Williams & W Pulawski, AMS, 1 worker; 29k S Tennant Creek, 20 May 1973, CA&TF Houston, WAM, 1 worker, 1 male; 6k NE Barkly Roadhouse, 7 Aug 2012, M Batley, AMS, 1 worker; Entire Ck, 155k NE byE Alice Springs, 6 Oct 1975, JC Cardale, ANIC, 1 worker; 53k E byN Alice Springs, 6 Oct 1975, JC Cardale, ANIC, 4 workers; 34k E Alice Springs, 7 Oct 1995, K Walker, NMV, 1 worker; Simpsons Gap, 4 Mar 1995, K Walker, 2 workers, 7 Oct 1995, K Walker, NMV, 1 worker; 45k SW Alice Springs, 24–25 Sep 1996, AED & LJD, CS-11: 20 workers, CS- 13: 21 workers; 89k W Alice Springs, 1k W Ellery Big Hole, 10 Oct 1995, K Walker, NMV, 1 worker; 91k W Alice Springs, 3k W Ellery Big Hole, 10 Oct 1995, K Walker, NMV, 1 worker; 95k W Alice Springs, 10 Oct 1995, K Walker, NMV, 1 worker; Serpentine Gorge, 7 Oct 1994, K Walker, NMV, 3 workers; Ormiston Gorge, 24 Feb 1994, K Walker, NMV, 12 workers; Ormiston Red Gum site, 10 Oct 1995, K Walker, NMV,> 20 workers; 50k along Haasts Bluff Road (ca. 180k W Alice Springs), 11 Oct 1995, K Walker, NMV, 15 workers. Western Australia: Samples collected from flowers or traps: Wyndham, 17 Nov 1930, TG Campbell, ANIC, 2 workers; 45k NE Halls Creek on Great Nth Hwy, 17 Apr 1980, G Anderson & D Symon, WAM, 5 workers; Moola Bulla Station, 1 Oct 1994, R Patterson, WAM, 12 workers; 50k SW Halls Creek on Great Nth Hwy, 6 Apr 1980, G Anderson & D Symon, WAM, 6 workers.

Austroplebeia australis –Western Colour Morph. Western Australia: Samples collected from flowers, water pools or traps: 12k S Kalumburu, CALM Site 13/4, 7–11 Jun 1988, TA Weir, ANIC, 3 workers; 50k S Kalumburu, 14 Sep 1985, AED & LJD, WS-11: 10 workers; Walsh Point, 17 May 1983, ID Naumann & JC Cardale, ANIC, 4 workers; 1k W Walsh Point, 10 May 1983, ID Naumann & JC Cardale, ANIC, 4 workers; Mitchell Plateau, Lone Dingo, 9–19 May 1983, ID Naumann & JC Cardale, ANIC, 1 worker; Mitchell Plateau, Mining Camp, 9–19 May 1983, ID Naumann & JC Cardale, ANIC, 6 workers; 4k S by W Mining Camp, 13 May 1983, ID Naumann & JC Cardale, ANIC, 1 worker; Carson Escarpment, 9–15 Aug 1975, IFB Common & MS Upton, ANIC, 2 workers; 70k N Drysdale River station, 18 Sep 1985, AED & LJD, WS-13: 2 workers; 45k N Drysdale River station, 14 Sep 1985, AED & LJD, WS-10: 9 workers; 30k N Drysdale River station, 12 Sep 1985, AED & LJD, WS-9: 2 workers; Drysdale River station, 12–18 Sep 1985, AED & LJD, WS-8: 2 workers, WS-14: 9 workers; 25k N Gibb River Station, 12 Sep 1985, AED & LJD, WS-7: 4 workers.

Austroplebeia australis: Nest and flower samples with mixed characteristics. Queensland: Nest samples, collected by AED & LJD except where otherwise stated: 110k S Innot Hot Springs, 6 Oct 1998 (Nest M33, 14 workers, 2 males); Duaringa, 15 Nov 1987 (Nest N30, 7 workers); Willows, 14 Sep 1996 (Nest C45, 14 workers); Taroom, Oct 2014, A Beil, (Nest D60, 17 workers). — Samples collected from flowers or traps: 13k SE Coen, 1 Nov 1988, K Walker, NMV, 2 workers; Silver Plains, 55k SE Coen, 1 Nov 1988, K Walker, NMV, 2 workers; Artemis Station, 28 Sep 1998, AED & LJD, MS-16: 4 workers; Palmer River on Mulligan Hwy, 18 Feb 1982, AED & LJD, QS-8: 4 workers; 10k S Bowen, 14 Feb 1982, AED & LJD, QS-5: 16 workers; 11K SSW Rolleston, 15 Apr 2005, M Batley, AMS, 2 workers; 45k W Mitchell, 14 Apr 2005, M Batley, AMS, 3 workers; 12k SW Charleville, 28 Aug 2009, M Batley, AMS, 1 worker; 48k WSW Charleville, 28 Aug 2009, M Batley, AMS, 1 worker; 50k S St George, 14 Sep 2005, M Batley, AMS, 4 workers; 25k S Dirranbandi, 15 Sep 2005, M Batley, AMS, 1 worker. New South Wales: Samples collected from flowers or traps: Lightning Ridge, 15 Sep 2005, M Batley, AMS, 1 worker; Coonabarabran, 2 Nov 1968, TF Houston, WAM, 1 male; 32k E Cobar, 2 Oct 2005, M Batley, AMS, 1 worker. Northern Territory: Samples collected from flowers or traps: Cooper Creek, 11k S byW Nimbuwah Rock (ca. 30k NE Oenpelli), 3–4 Jun 1973, JC Cardale, ANIC, 1 worker; ca. 60k E Alice Springs, 22 Sep 1996, AED & LJD, CS-8:> 60 workers. Western Australia: Samples collected from flowers or traps: Phillips Range, 12 May 1973, CA&TF Houston, WAM, 3 workers; King Leopold Ranges, 4k NW Rifle Point, 12 Apr 1980, G Anderson & D Symon, WAM, 3 workers; 1k NE Mt Bell, 28 Jul 1988, TF Houston, WAM, 12 workers; Durack River, Jack's Hole, 19 Sep 1985, AED & LJD, WS-15: 1 worker.

Queen specimens examined.

Austroplebeia australis –Eastern Colour Morph. Nest samples collected by AED & LJD except where otherwise stated, 18 queens: Queensland: Cape York near Bamaga, 10 Aug 1983 (Nest F31, 1 virgin); Miles, Nov 2002, Robert Raymond, 1 gravid; Tara, Nov 2013, Allan Beil, 8 gravid, Jun 2015, Allan Beil, 5 gravid. New South Wales: Tenterfield, Dec 1980, 1 gravid; Kempsey, Jan 1980 (Nest 15, 1 gravid), Mar 1980, 1 gravid.

Austroplebeia australis –Central Colour Morph. Nest samples collected by AED & LJD, 9 queens: Queensland: Croydon, 31 Aug 1987 (Nest N21, 1 gravid), 19–20 Aug 1990 (Nest G1, 4 virgin); Hughenden, 30– 31 Aug 1983, (Nest F53, 1 gravid); Northern Territory: 50k W Timber Creek, 27 Sep 1985 (Nest W13, 1 gravid; Nest W17, 2 virgin).

Remarks. The type locality of the species A. australis is Central Australia, NT. However, even in the type description, Friese (1898) considered that the Central Australian bees resembled QLD specimens from Burnett River, near Bundaberg. Michener (1961) reinforced this, noting similarities between bees from Central Australia, NT, and Brisbane, QLD.

As the popularity of stingless beekeeping in QLD grew in the 1980s, the name A. australis became widely applied to the bees of the Eastern Colour Morph in QLD. However, data from this study indicate that the lectotype of A. australis actually belongs to the Central Colour Morph population.

The A. australis lectotype. Historical sources (explained above) suggest that lectotype specimen of A. australis was probably collected in the Hermannsburg area of Central Australia, which is well within the distribution of the Central Colour Morph ( Fig. 28 c). Furthermore, it is closest to the Central Colour Morph in HTW/HBW ( Fig. 11 , Table 3 ). However, its body colouration is darker (Thorax intermediate between Grades 4 and 5, Figs 5 , 7 , 10 ), resembling that of the Eastern Colour Morph.

Some other bees with darker colouration (Thorax Grade 4), caught on flowers in the MacDonnell Ranges, were found in the NMV and in our collection. Their localities were only 40 to 160 km from Hermannsburg, NT. Sting rudiment analysis of two of these bees ( Fig. 15 ) showed that they belong to the Central Colour Morph despite their darker colouration. So the Central Colour Morph of Central Australia does include some bees with darker colouration. We conclude that the A. australis lectotype probably belongs to the Central Colour Morph.

We consider that the primary types of A. percincta , A. cockerelli , A. ornata and A. websteri also all belong to the Central Colour Morph (see below). However, as A. australis is the earliest described name, we conclude that A. australis is, by priority, the correct name for the Central Colour Morph populations of NT and western QLD.

We consider the Eastern and Central Colour Morphs to be conspecific (as explained in 'Species determinations for the groups of Halcroft et al. (2015) ' in the Results section, above). So the name A. australis should still be used for Eastern Colour Morph colonies in QLD. However, if future studies demonstrate species differences between the Eastern and Central Colour Morphs, a new species name will need to be applied to the Eastern Colour Morph colonies.

The A. percincta lectotype. This specimen is consistent with the Central Colour Morph of A. australis in most characteristics. It has a Face Colour Grade of 4 and a Thorax Colour Grade of 12 ( Figs 1 , 3 , 5 , 7 , 10 ). However, its HTW/HBW score is borderline between those of the Central and Eastern Colour Morphs ( Fig. 11 , Table 3 ). Its type locality (Hermannsburg, NT) is well within the geographic range of the Central Colour Morph ( Fig. 28 c). We conclude that the A. percincta lectotype belongs to the Central Colour Morph.

The holotypes of A. cockerelli and A. websteri . These specimens clearly match the Central Colour Morph of A. australis in both structure ( Fig. 11 , Table 3 ) and colouration ( Figs 1 , 3 , 5 , 7 , 10 ). Their faces are discoloured but are probably Grade 4, and their Thorax Colour Grade is 11. Their type localities (Borroloola, NT, and Wyndham, WA respectively) were also well within the geographic range of the Central Colour Morph ( Fig. 28 c).

The A. ornata type series. Comparison of the A. ornata type specimens from Cape York, QLD, with recently collected bees was hampered by their poor condition: collapsed eyes, discolouration, missing legs and surface contamination. However, the lectotype, and two of the paralectotypes (HYM-34696, HYM-34697) match A. australis –Central Colour Morph in most characteristics. In the lectotype, the face is Grade 4 and thorax is probably Grade 11 ( Figs 1 , 3 , 5 , 7 ). In the type description, the colour markings on the face, thorax and apical terga given for the worker are also consistent with those of A. australis –Central Colour Morph. Its HTW/HBW score is borderline between those of the Central and Eastern Colour Morphs ( Fig. 11 , Table 3 ). Its eyes are collapsed, preventing a meaningful head width measurement. In the two paralectotypes, the colouration is consistent with that of A. australis –Central Colour Morph (both specimens: Face Grade 4, Thorax Grade 11, Figs 1 , 3 , 5 , 7 ). HTW/HBW is not measureable in HYM-34696, but in HYM-34697 these scores are in the Eastern Colour Morph range. Geographically, the A. ornata type series was collected from a region of Cape York where the Central Colour Morph transitions towards the Eastern Colour Morph ( Fig. 28 c).

Two paralectotypes (HYM-34695 and one ANIC worker) correspond with A. cassiae in colouring (Thorax Grades 0 and 2, Figs 5 , 7 ), pilosity and structure (HBW of ANIC specimen 0.44 mm), where these characters are scorable. This explains the comment in Rayment's type description, 'On some specimens from the same “nest” the yellow of the worker's scutellum is limited to two dots' ( Rayment 1935). This specimen series, collected by Thomson, is actually a mixture of two species, so they would not have come from a single nest. However, in central-western Cape York, AED & LJD found nests of A. australis –Central Colour Morph and A. cassiae in close proximity: in the same tree near Kowanyama (nests G8 and G9) and within 25m at Koolatah Station (nests M17 and M22).

In the ANIC male paralectotype, gonostylus shape and pilosity characters show that it does not belong to A. essingtoni nor A. cincta . However, its species could not be further identified. The other ANIC worker paralectotype belongs to either the Central or Eastern Colour Morph of A. australis . This specimen only has a blue 'cotype' label, but has no other label giving species name, collection locality or collector. Unlike the other specimens, its eyes are not collapsed. It is unclear whether this specimen belongs to Thomson's Cape York specimen series.

Individuals with orange abdomens. Some workers and males from both the Central and Eastern Colour Morphs had distinctive orange abdomens ( Fig. 26 ), particularly vivid in colour when viewed against bright light. These colonies were distributed from Cape York, QLD to coastal NSW, but they were particularly common in western QLD. In some colonies this colouring is present in all individuals, whilst in others some bees may have brown or black abdomens. We consider this characteristic to be a simple colour variant within A. australis .

......continued on the next page *Variables: As given in Table 2.

Other than the types, the following worker specimens were measured. A. australis –Eastern Colour Morph, N=21 unless otherwise stated, from nests C32 (4x), F31 (4x), NSW15 (4x), S9 (5x), S31 (4x). A. australis –Central Colour Morph, N=16 unless otherwise stated, from nests C49 (4x), F53 (4x), M17 (4x), W17 (4x). A. australis –Western Colour Morph, N=5 unless otherwise stated, from WA flower samples WS-11 (1x), Walsh Point (2x), Mining Camp (2x). A. N=20; B. N=15; C. 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 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 5. Thorax colour marking grades used to classify workers and males of Austroplebeia.

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 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 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 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 26. An A. australis worker with the distinctive orange abdomen, that is common in western QLD populations (photograph by D. Clarke).

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.

Tables

TABLE 3. Structural dimensions from workers of A. australis. Measurements in mm. A " – " indicates that measurements were not available.

australis Lectotype. A VAR *	cockerelli Holotype. A	ornata Lectotype. A percincta Lectotype. A websteri Holotype. A	percincta Lectotype. A websteri Holotype. A	websteri Holotype. A	A. australis Eastern Colour Morph, other workers Mean SD	Central Colour Morph, other workers Mean SD	Western Colour Morph, other workers Mean SD
1 4.20	3.85	4.35	4.25	4.66	3.94 0.21	3.92 0.30	4.31 0.37
2 1.94	1.78	–	1.90	1.70	1.78 0.05	1.73 0.04	1.83 0.03
3 1.56	1.56	1.44	1.62	1.48	1.48 0.04	1.43 0.03	1.51 0.01
4 1.26	1.12	1.14	1.30	1.14	1.17 0.03	1.15 0.04	1.21 0.02
5 1.20	1.10	1.16	1.20	1.10	1.13 0.05	1.09 0.03	1.12 0.02
6 1.32	1.18	1.16	1.30	1.18	1.22 0.06	1.17 0.02	1.22 0.02
7 1.00	0.94	0.92	0.90	0.92	0.95 0.04	0.93 0.03	0.98 0.01
8 0.09	0.15	0.17	0.11	0.16	0.16 0.01	0.16 0.01	0.17 0.01
9 0.44	0.43	0.42	0.48	0.41	0.42 0.02	0.42 0.02	0.45 0.01