Heteropus variegatus Macleay, 1877 Proc. Linn. Soc. N.S.W . Heteropus maculatus de Vis. Proc. Roy. Soc. Qld Systematics of the Carlia ‘ fusca’ complex (Reptilia: Scincidae) from northern Australia Donnellan, S. C. Couper, P. J. Saint, K. M. Wheaton, L. Zootaxa 2009 2009-09-10 2227 1 1 31 (Macleay, 1877) Macleay 1877 [151,566,151,177] Reptilia Scincidae Carlia Animalia Squamata 17 18 Chordata species sexdentata   Figs 9, 10, & 11: Tables 3, 4& 5  1877  Heteropus sexdentatusMacleay. Proc. Linn. Soc. N.S.W.2: 67. Cape Grenville, north-eastern Queensland. LectotypeAMS R31879.   1877  Heteropus variegatus Macleay, 1877 Proc. Linn. Soc. N.S.W.2: 66.  Darnley Island, N Queensland. LectotypeAMS R31868.  1885  Heteropus maculatusde Vis. Proc. Roy. Soc. Qld1: 169. Cape York, north-eastern Queensland. Typematerial missing, see Covacevich (1971).   1885  Heteropus rubricatusde Vis. Ibid.1: 170. Cape York, north-eastern Queensland. Typematerial missing, see Covacevich (1971).  Material examined:   Lectotype:AMS R31879. Cape Grenville, north-eastern Queensland( 11°58'S, 143°14'E).  Other material: Queensland. Torres strait -(Vouchers with associated genetic samples):  QM J86364, Saibai Island( 9°24'S, 142°41'E);  QM J86350, Dauan Island( 9°25'S, 142°32'E);  QM J86353,  QM J86363, Masig, Yorke Island( 9°45'S, 143°24'E);  QM  J86360-2, Yam Island( 9°54'S, 142°46'E). Cape York Peninsula -  QM J68594, Cape York, Somerset( 10°44'3”S, 142°35'3”E);  QM J45598,  QM  J45601-2,  QM J45599, near Telegraphcrossing, north bank of Jardine River, ( 11°08'S, 142°22'E);  QM J26200, Heathlands, site 5 ( 11°39'S, 142°50'E);  QM J26216,  QM J26220,  QM J26224 Cape York, Heathlands( 11°39'S, 142°50'E);  QM J39745 7.8kmN mouth of Janie Creek( 11°57'S, 141°52'E);  QM J46366, near Harmer Creek,  19kmE Nixon’s Homestead, Shelburne Station( 11°58'S, 142°55' E);  QM J46341, 21.8kmfrom Nixon’s Homestead, on road to Double Point( 11°58'S, 142°05'E);  QM J46333, Harmer Creek,  18kmE Nixon Homestead, Shelburne Station( 11°59'S, 142°53'E);  QM J67615, Haggerstone Island( 12°03'S, 143°18'E);  QM J80416, Pennyfather River,  70kmN Weipa( 12°15'39”S, 141°42'40”E);  QM J74714, Thong Treesouth of Pennyfather River( 12°19'23"S, 141°41'26"E);  QM J32017, Kungathan( 12°22' S, 143°12'E);  QM  J86997-8, Glennie Tableland( 12°27'43"S, 142°55'26E);  QM J62356, Weipa area( 12°28'41"S, 141°50'37”E); QMJ62357, Weipa area( 12°28'01"S, 141°50'34”E);  QM J70190, Weipa area( 12°28'37"S, 141°50'03”E);  QM  J86984-7 Pascoe Rivermouth ( 12°29'09"S, 143°16'28"E);  QM J62354, Weipa Area( 12°30'28"S, 141°47'52"E);  QM  J34394-5 Portland Roadsturnoff, Iron Range( 12°38'S, 143°23'E);  QM J70340, Weipa area( 12°43'22"S, 141°56'48"E);  QM J34389, W Claudiecampsite, Iron Range, ( 12°46'S, 143°17'E);  QM J39716,  QM J39748, 6km SW Boyd Point, Pera Head( 12°57'S, 141°36'E);  QM J34478, Buthen Buthen, Nesbit River, E Coen( 13°21'S, 143°27'E);  QM  J41927-9, Thanmala, via Aurukun( 13°23'S, 141°42'E);  QM J34502,  QM J34505/7, Steen’s Hut, approx.  30kmNE Coen( 13°34'S, 143°13'E);  QM  J86893-4, Kulla NP, McIlwraith Range( 13°42'36”S, 143°18'36”E);  QM J37487, 12km NW Coen( 13°51'17"S, 143°9'1"E);  QM  J46240-1,  QM  J46245-6, McIlwraith Range( 13°50'S, 143°17'E);  QM J78269, 5km N Coen( 13°54'08”S, 143°10'56”E);  QM J78264, adjacent to ranger station Coen Creek, Coen( 13°56'2”S, 143°11'59"E);  QM J78341approx. 3kmS Coen( 13°57'44”S, 143°11'24”E);  QM J78266S Coen, ( 14°08'S, 143°13'31”E);  QM J60337, Stanley Island, Flinders Group NP( 14°8'58"S, 144°15'15”E);  QM J72702, Lilyvale Station( 14°25'5"S, 143°45'42"E).  Northern Territory -(all specimens from NTMunless otherwise stated): R19025, Cape Wessel Island( 11°S, 136°46'E);  R26299,  R26310, Jensen’s Bay, Marchinbar Island( 11°10'S, 136°41'E);  R08950, S Nip Point Marchinbar Island( 11°21'S, 136°33'E);  R28607-8, Rimbija Island, Wessel Islands( 11°34'S, 136°45'E);  R22757,  R22761/5, Astell Island, English Company Isles( 11°52'S, 136°25'E);  QM J84006 Maningrida( 12°02'52”S, 134°13'20”E);  QM  J86281-2, Maningrida( 12°02'55”S, 134°13'33”E);  QM  J86284-5,  QM J86287, ManingridaCEC ( 12°03'08”S, 134°13'20”E);  R23948, Ramingining area, Arafura Swamp( 12°10'08”S, 134°58'04”E);  R23945, Ramingining area, Arafura Swamp( 12°12'S, 134°59'E);  R16156/8,  R16161-2, Blyth Rivercrossing, Arnhem Land( 12°21'S, 134°41'E);  R23902, Djapididjapin Creek, near Ramingining, Arafura Swamp( 12°21'50”S, 134°54'19”E);  R23959, Ramingining Area, Arafura Swamp(12° 12°30S, 134°58’56”E);  R23995, Ramingining Aarea, Arafura Swamp( 12°12'25”S, 134°59'11”E);  R28280, Groote Eylandt,  Gulfof Carpentaria( 13°56'S, 136°36'E);  R03357, Angurugu, Groote Eylandt( 13°58'S, 136°26'E);  R07566, Angurugu, Groote Eylandt( 13°59'S, 136°28'E).   Distribution:North-eastern Northern Territoryand north-eastern Queensland. Northern Territory– northeastern Arnhem Land, Crocodile Island, The Wessel Islands, the English Company’s Islands and Groote Eylandt. Queensland- Torres Strait Islands and Cape York Peninsula from the head of Princess Charlotte Bay ( 14°30'S) north.   Diagnosis:  Carlia sexdentatamost closely resembles  C. longipesfrom which it is separated herein. The two species are distinguished most readily by the colour pattern of adult males and the nature of the ear lobules. In  C. sexdentatathe dark midlateral zone between ear and shoulder does not contrast sharply with the lower neck colour vscontrasting sharply with lower neck colour; in  C. sexdentatathe ear aperture usually has a well-developed series of lobules on the anterior margin often producing a distinctive comb- like effect, lobules poorly to moderately developed on other surfaces of ear ( Fig. 10a, b) vsear aperture completely surrounded by sharply pointed lobules ( Fig. 7).  Carlia sexdentatalacks the dorsal and lateral pattern seen in  C. quinquecarinata(large, longitudinally aligned, dark-edged, pale dashes on the dorsum and a broken midlateral line [ Fig. 12c]).  Carlia sexdentataneed only be distinguished from other species of  Carliawith a smoothly rounded posterior edge to the midbody scales. It is distinguished from  C. mundaand  C. tetradactylaby the nature of its ear lobules (sharply pointed vssmall and rounded); from  C. rimulaby its greater midbody scale count and larger size (31–38 vs≤ 30; max SVL = 63.5mm vs 39mm); from  C. rhomboidalisand  C. rubrigularisby the state of the interparietal scale (free vsfused); from  C. rostralisby its ear lobules and male breeding colours (ear aperture usually has a series of well-developed lobules on the anterior margin often producing a distinctive comb-like effect, lobules poorly to moderately developed on other surfaces of ear vsear aperture generally with only one or two large pointed lobules on anterior edge; males with pale throat vsblack throat). Of the above listed species, its broad distribution only overlaps with  C. mundaand  C. rimula.Whether it overlaps with  C. longipesremains unclear. The two species may be mutually exclusive but, if so, the southern limit for  C. sexdentatais in close proximity to the northern limit for  C. longipes.   Description:SVL (mm) 21.00–63.51, n = 83. Proportions as %SVL(mean ± standard deviation): TL 170.23–198.18 (185.37 ± 7.53, n = 20); AG 41.58–54.58 (48.19 ± 2.70, n = 82); L1 28.81–42.03 (34.69 ± 2.41, n = 82); L2 42.11–58.51 (50.84 ± 3.88, n = 82); HL 20.14–25.42 (22.75 ± 1.08, n = 81) Eye-ear 6.86–9.10 (8.01± 0.47, n = 81); Snout 7.86–10.74 (9.16± 0.53, n = 81). Body robust. Head barely distinct from neck. HW 59.15-79.30% HL (68.28 ± 3.44, n = 81). Limbs moderate. L1 59.42–80.42% L2 (68.40± 4.11, n = 82).  Scalation:Rostral in broad contact with frontonasal. Prefrontals large, narrowly (42% of specimens), moderately (51% of specimens) or widely (6% of specimens) separated; contacting in QM J86281. Supraoculars 4; 1 and 2 incontact with frontal, 2, 3 and 4 incontact with frontoparietal. Frontoparietals fused, forming a single shield. Interparietal free. Enlarged nuchal scales 2–3 (mode = 2, n = 862). Snout rounded in profile. Loreals 2 ( 1 inQM J86284). Preoculars 2. Presubocular single. Supraciliaries 6–8 (7.05±0.31, mode = 7, n = 82). Lower eyelid moveable with clear window; palpebral disc small, occupying half or less of lower eyelid. Ear aperture subequal or larger than palpebral disc; usually vertically aligned (68% of specimens) but sometimes round (32% of specimens), with an enlarged, comb-like series of sharp lobules on anterior margin, usually some elongated sharp lobules projecting down from the posterior edge of the dorsal margin (66% of specimens) and sometimes a series of smaller lobules along the posterior margin (30% of specimens). Supralabials 6–8 (7.01±0.19, mode = 7, n = 82), with fourth to sixth below centre of eye (5.01±0.19, mode 5, n = 82). Infralabials 5–8 (mode = 6, n = 83). Dorsal scalation smooth to weakly tricarinate; posterior edge smoothly curved. Midbody scale rows 31–38 (mean = 34.64, mode = 34, SD = 1.62 n= 81). Paravertebral scale rows 44–50 (mean = 47.09 mode = 47, SD = 1.67, n = 79). Lamellae beneath 3 rdfinger 17–25 (21.11±1.55, mode = 21, n = 82). Lamellae beneath 4 thtoe 24-37 (30.61±2.44, mode = 32, n = 82). The scale counts and body proportions of the Queenslandand Northern Territory  C. sexdentatapopulations are broadly overlapping (see Table 5).  Colour pattern in spirit:Dark olive brown dorsally; uniform, or with dark streaks aligned longitudinally on scale edges. Tail usually with a longitudinal series of dark-edged pale flecks. Venter off-white.  Colour pattern in life: adult males in breeding colours( Fig. 9a, d & f): usually with some indication of a poorly defined, white dorsolateral stripe (commencing behind supraciliaries, running through upper secondary temporals and terminating just behind forelimb). This is bordered below by a dark upper lateral zone (a grey to blackish smudge, encompassing 2–3 scales rows and extending from eye to just behind forelimb) which does not contrast sharply with the scales of the lower neck which, though paler, are often dark-edged. Dark upper lateral zone terminates just behind the forelimb and is continuous with the metallic pink flank colouration (rich orange in life). Limbs and sides of tail orange- brown. Females( Fig. 9b, c): with a narrow, pale facial stripe that begins beneath the eye and terminates near the dorsal margin of the ear aperture. The dark upper lateral is enclosed between narrow, white dorsolateral and midlateral stripes which are most prominent anterior to the forelimb (the former commencing behind the eye, the latter behind the ear). This pattern extends varying distances along the flanks (sometimes approaching the hindlimb insertion). The upper lateral zone is broken by one or more, obscure vertical, white bars in front of the forelimb and has irregular white dashes along its full length. Dorsum often with white flecking and sometimes a dark vertebral streak ( Fig. 9b). Juveniles:similar to females but with more prominent pale flecking on dorsum and flanks. NT populations( Fig. 9e): general pattern similar to Queenslandmaterial, but less prominent. Dark upper lateral zone usually vague and sometimes broken into a series of narrow, longitudinal stripes.  Measurements and scale counts of Macleay  Heteropus sexdentatus types:AMS R31879-81: SVL 38.43–53.59mm, n = 3; Proportions as %SVL (mean ± standard deviation): TL 200.03 (n = 1); AG 45.49– 51.83 (48.80 ± 3.18, n = 3); L1 34.63–38.10 (36.52 ± 1.74, n = 3); L2 54.0–58.05 (55.53 ± 2.20, n = 3); HL 23.31–23.80 (23.61 ± 0.27, n = 3); Eye – ear 8.12–8.60 (8.35 ± 0.24, n = 3); Snout 9.60–10.67(10.19 ± 0.55, n = 3) Body robust. Head barely distinct from neck. HW 60.87–71.58% HL (65.08 ± 5.71, n = 3). Limbs moderate. L1 64.18–67.52% L2 (65.76 ± 1.68, n = 3).  Raw measurements and scale counts of lectotype( Fig. 11a): AMS R31879. SVL 53.59 mm; AG 26.25 mm; L1 18.56 mm; L2 28.92 mm; HL 12.49 mm; HW 8.94 mm; Eye – ear 4.61; Snout 5.14: midbody scale rows 36; paravertebral scale rows 48; supraciliaries 7; supralabials 7; fifth supralabial below eye; infralabials 6; subdigital lamellae beneath 3 rdfinger 23; subdigital lamellae beneath 4 thtoe 34; enlarged nuchals 2; preoculars 2; presuboculars 1; postsupralabial divided; temporals – 1 primary, 2 secondary; ear vertical with sharp lobules at front and top – those along dorsal margin small and deeply recessed ( Fig. 11b); palpebral disc smaller than ear opening. The scales of the dorsal and lateral surfaces are weakly tricarinate.   FIGURE 11.(a) AMS R31879, Lectotype of  H. sexdentatus Macleay 1877Cape Grenville, north-eastern Queensland (image: J. Wright). (b) Ear lobules, left side of AMS R31879. Note that the lobules on the dorsal margin are small and deeply recessed, hence not clearly discernible. (image: G. Thompson). (c) AMS R31868, Lectotype of  H. variegatus Macleay 1877DarnleyIsland, north-eastern Torres Strait (image: J. Wright). (d) Ear lobules, left side of AMS R31868. Note that the lobules on the dorsal margin are small and deeply recessed, hence not clearly discernible. (image: G. Thompson). Pattern of   lectotype:AMS R31879 is badly faded and has lost all traces of patterning, as has AMS R31881. The  paralectotype, AMS R31880, still shows traces of the typical female colour pattern, i.e. a dark upper lateral zone bordered by a pale dorsolateral and midlateral stripe; broken by obscure, vertical, white bars in front of forelimb.   Comments:The diagnostic characters provided herein, separating  C. longipesfrom  C. sexdentatahold well for mainland  C. sexdentatapopulations. However, the ear lobule state (ear aperture is completely surrounded by sharply pointed lobules in  C. longipesvsusually a series of well-developed lobules on anterior margin often producing a distinctive comb-like effect, lobules poorly to moderately developed on other surfaces of ear for  C. sexdentata) is more variable in Torres Strait with island  C. sexdentataspecimens commonly approaching the`completely surrounded’  C. longipescondition. In Torres Strait, where the condition of the ear lobules may prove ambiguous, the diagnostic pattern differences hold well for adult males. Two specimens listed in material examined were not included in the statistical analysis (QM J67615, Haggerstone Island, and QM J72702, Lilyvale Station) as they represent outliers to the main distribution and have pattern and ear lobule characteristics that, although generally consistent with  C. sexdentata,are unusual. Both specimens have a single large lobule on the anterior margin of the ear (vs a ‘comb’ of enlarged lobules) and, although being females, exhibit a male-like colour pattern (anterior midlateral zone reduced to a dark smudge between ear and forelimb). Despite these inconsistencies, both specimens can be readily separated from  C. longipes. An issue arises concerning the names  C. sexdentataand  C. variegataand which of these is the senior synonym. Wells and Wellington (1985) resurrected  C. variegata( Macleay 1877)from the synonymy of  C. longipesas the appropriate name for the Torres Strait island ‘  C. fusca’, yet provided no explanation for doing so and made no mention of  H. sexdentatus Macleay 1877, which Macleay had published in the same paper. The action of Wells and Wellington (1985) potentially gives  C. variegatapriority over  C. sexdentata. However, while the lectotypesof  H. sexdentatusand  H. variegatusare faded both retain the key diagnostic pattern elements, but the lectotypeof  H. sexdentatushas the majority ear lobule condition ( Fig.11b) while the lectotypeof  H. variegatushas the minority condition for this trait ( Fig. 11d). Therefore we can associate more strongly the lectotypeof  H. sexdentatuswith our taxon  C. sexdentata. Furthermore, as this is the first time in which these species alone have been considered as competing synonyms rather than as junior synonyms of a third species,  C. longipes, we invoke the principle of the first reviser (ICZN, Article 24.2) and consider  H. sexdentatusas the more appropriately applied name, with  H. variegatusrelegated to the synonymy of  C. sexdentata. The provenance and affiliations of two other types,  Heteropus maculatusde Vis 1885and  H. rubricatusde Vis 1885need to be considered to complete our analysis of the synonymy of the ‘  C. fusca’ group in northern Australia. The descriptions of  Heteropus maculatusand  H. rubricatusare based on material collected on ‘Cape York’ by Kendall Broadbent some time prior to November 1884; the date that de Vis presented his ‘…conspect of the genus  Heteropus’ at a meeting of the Royal Society of Queensland(de Vis, 1885). Based on locality, colour pattern and scalation characters, these taxa clearly belong to the ‘  Carlia fusca’ group and accordingly were assigned to the synonymy of  C. longipesby Ingram and Covacevich (1989). With our resurrection of  C. sexdentata(new combination) from the synonymy of  C. longipes, it is necessary to revisit de Vis’ descriptions of  H. maculatusand  H. rubricatusto assess their taxonomic status. In the absence of specimens ( typesmissing, see Covacevich, 1971), the only clues to their identities lie in: 1) information on the whereabouts of the collector in the period preceding publication. Importantly, where on Cape York Kendell Broadbent did collect in the period leading up to de Vis’  Heteropusdescriptions? If he was north of Princess Charlotte Bay then these specimens are likely to belong to  C. sexdentata, if he was south,  C. longipesis the most likely identification; and 2) de Vis’ scant typedescriptions. 1) The report of the Board of Trustees for the QueenslandMuseum for the year 1884 states that at the start of the year Broadbent was ‘…dispatched to Somerset, Cape York, and succeeded in making a large collection chiefly of fish, but including novelties in each department of vertebrate life.’ (Anon, 1885). Broadbent then proceeded to the Kimberley region of Western Australiawhere he met with limited collecting success and subsequently revisited Somerset on his return trip home. Here he supplemented his previous collections and was back in Brisbane in September, 1884. A summary of his Cape York (= Somerset) collections ( Anon, 1985, Appendix iv) shows that 54 reptile specimens representing 23 species were collected on the first visit with additional specimens obtained on the return visit (numbers not specified in report). This information is further corroborated by collection data contained in the QueenslandMuseum’s vertebrate registers. 2) Both descriptions refer to ‘free scales’ along the fore-edge ( 5-6 in  H. maculatus, 3 in  H. rubricatus) of the ear aperture with no mention of lobules on the other edges of the ear, the typical  C. sexdentatastate. Further, the description of  H. maculatusprovides colour details that are consistent with male  C. sexdentatabut not  C. longipes(i.e.  H. maculatushas a broad lateral stripe, but no mention is made of this being bordered above by a distinct, pale dorsolateral). The lateral stripe of  H. rubricatusis less informative, being ‘…edged above and below with white…’ (de Vis, 1885). This pattern could apply to either  C. sexdentataor  C. longipes, but would suggest the specimen examined by de Vis was likely to have been a female. The other descriptive details, i.e. measurements and midbody scale counts, fall within the range of both species. On balance, the evidence points to the missing de Vis typesbeing referable to  C. sexdentata. It is likely that these specimens were collected at Somerset on northern Cape York Peninsula ( 10°45'S) in 1884. The ear lobule descriptions for both de Vis taxa point to an identification of  C. sexdentataas does the colour pattern description of  H. maculatus. It is likely that both  H. maculatusand  H. rubricatusare conspecific with  C. sexdentata, the former description depicting a male, the latter a female.  Heteropus maculatusand  H. rubricatusare, herein, formally assigned to the synonymy of  C. sexdentata. R31879 R Cape Grenville 17 18 1 Queensland lectotype R31868 R Darnley Island, N Darnley Island 17 18 1 Queensland lectotype Cape York 17 18 1 Queensland Cape York 17 18 1 Queensland R31879 R -11.966666 Cape Grenville 1294 143.23334 17 18 1 Queensland lectotype J86364 QM -9.4 Saibai Island 1300 142.68333 17 18 1 Queensland J86350 [558,1114,689,715] QM -9.416667 Dauan Island 1300 142.53334 17 18 1 Queensland J86353 [1127,1272,689,715] QM 17 18 1 Queensland J86363 QM Masig -9.75 Yorke Island 1299 143.4 17 18 1 Queensland J86360 [633,1177,727,753] QM -9.9 Yam Island 1298 142.76666 17 18 J86360-2 1 Queensland J68594 [169,879,766,792] QM Cape York & Somerset -10.734167 21 142.58417 17 18 1 Queensland J45598 [892,1036,766,792] QM 17 18 1 Queensland J45601 [1049,1217,766,792] QM 17 18 J45601-2 1 Queensland J45599 QM Telegraph -11.133333 Jardine River 1296 142.36667 17 18 1 Queensland J26200 QM Heathlands -11.65 1295 142.83333 17 18 1 Queensland J26216 [400,543,844,870] QM 17 18 1 Queensland J26220 [555,698,844,870] QM 17 18 1 Queensland J26224 [711,1376,844,870] QM Cape York & Heathlands -11.65 1295 142.83333 17 18 1 Queensland J39745 QM -11.95 Janie Creek 1294 141.86667 17 18 1 Queensland J46366 QM -11.966666 Shelburne Station 1294 142.91667 Harmer Creek 17 18 1 Queensland J46341 QM Double Point -11.966666 Nixon's Homestead 1294 142.08333 17 18 1 Queensland J46333 QM -11.983334 Shelburne Station 1294 142.88333 Harmer Creek 17 18 1 Queensland J67615 [493,1124,1000,1026] QM -12.05 Haggerstone Island 1294 143.3 17 18 1 Queensland J80416 QM -12.260834 Pennyfather River 21 141.7111 17 18 1 Queensland J74714 QM Thong Tree -12.323055 Pennyfather River 21 141.69055 17 18 1 Queensland J32017 [486,1026,1078,1104] QM Kungathan -12.366667 1293 143.2 17 18 1 Queensland J86997 QM Glennie Tableland -12.461945 21 142.92389 17 18 J86997-8 1 Queensland J62356 [488,1131,1117,1143] QM -12.478055 Weipa area 21 141.84361 17 18 1 Queensland QMJ62357 -12.466944 Weipa area 21 141.84277 17 18 1 Queensland J70190 [484,1098,1156,1182] QM -12.476944 Weipa area 21 141.83417 17 18 1 Queensland J86984 QM -12.485833 Pascoe River 21 143.27444 17 18 J86984-7 1 Queensland J62354 [592,1249,1194,1220] QM -12.507778 Weipa Area 21 141.79779 17 18 1 Queensland J34394 QM Portland Roads & Iron Range -12.633333 1293 143.38333 17 18 J34394-5 1 Queensland J70340 [820,1431,1234,1260] QM -12.722777 Weipa area 21 141.94667 17 18 1 Queensland J34389 [151,940,1273,1299] QM W Claudie & Iron Range -12.766666 1292 143.28334 17 18 1 Queensland J39716 [953,1095,1273,1299] QM 17 18 1 Queensland J39748 QM SW Boyd Point & Pera Head -12.95 1291 141.6 17 18 6 1 Queensland J34478 QM Buthen Buthen -13.35 Coen 1291 143.45 Nesbit River 17 18 1 Queensland J41927 [286,1001,1350,1376] QM Thanmala & Aurukun -13.383333 1291 141.7 17 18 J41927-9 1 Queensland J34502 [1014,1158,1350,1376] QM 17 18 1 Queensland J34505 QM Steen's Hut -13.566667 30 km NE Coen 1290 143.21666 17 18 1 Queensland J86893 QM Kulla NP & McIlwraith Range -13.71 21 143.31 17 18 J86893-4 1 Queensland J37487 [343,1041,1428,1454] QM -13.854723 Coen 21 143.15027 17 18 12 1 Queensland J46240 [1058,1236,1428,1454] QM 17 18 J46240-1 1 Queensland J46245 QM McIlwraith Range -13.833333 1289 143.28334 17 18 J46245-6 1 Queensland J78269 [630,1274,1467,1493] QM -13.902222 Coen 21 143.18222 17 18 5 1 Queensland J78264 QM -13.933889 Coen 21 143.19972 Coen Creek 17 18 1 Queensland J78341 QM -13.962222 Coen 21 143.19 17 18 1 Queensland J78266 [508,1071,1545,1571] QM -14.133333 Coen 925 143.22527 17 18 1 Queensland J60337 QM Flinders Group NP -14.149445 Stanley Island 21 144.25417 17 18 1 Queensland J72702 [736,1431,1584,1610] QM -14.418056 Lilyvale Station 21 143.76167 17 18 1 Queensland R19025 NTM, R -11.0 Cape Wessel Island 0 136.76666 17 18 1 Northern Territory R26299 [290,393,1662,1688] R 17 18 1 Northern Territory R26310 [402,1166,1662,1688] R -11.166667 Marchinbar Island 1295 136.68333 Jensen's Bay 17 18 1 Northern Territory R08950 R -11.35 Nip Point Marchinbar Island 1295 136.55 17 18 1 Northern Territory R28607 [651,1430,1701,1727] R -11.566667 Wessel Islands 1295 136.75 Rimbija Island 17 18 R28607-8 1 Northern Territory R22757 [151,256,1740,1766] R 17 18 1 Northern Territory R22761 [264,1123,1740,1766] R -11.866667 English Company Isles 1294 136.41667 Astell Island 17 18 1 Northern Territory J84006 QM Maningrida -12.047778 21 134.22221 17 18 1 Northern Territory J86281 [505,1185,1778,1804] QM Maningrida -12.048612 21 134.22583 17 18 J86281-2 1 Northern Territory J86284 [1200,1374,1778,1804] QM 17 18 J86284-5 1 Northern Territory J86287 QM Maningrida -12.052222 21 134.22221 17 18 1 Northern Territory R23948 R -12.168889 Arafura Swamp 21 134.96777 Ramingining area 17 18 1 Northern Territory R23945 [500,1295,1857,1883] R -12.2 Arafura Swamp 1293 134.98334 Ramingining area 17 18 1 Northern Territory R16156 [1309,1431,1857,1883] R 17 18 1 Northern Territory R16161 [151,990,1895,1921] R Arnhem Land -12.35 Blyth River 1293 134.68333 17 18 R16161-2 1 Northern Territory R23902 R Djapididjapin Creek -12.36389 Arafura Swamp 21 134.90527 Ramingining 17 18 1 Northern Territory R23959 R -12.5 Arafura Swamp 925 134.98221 Ramingining Area 17 18 1 Northern Territory R23995 [505,1431,1973,1999] R -12.206944 Arafura Swamp 21 134.98639 Ramingining Aarea 17 18 1 Northern Territory R28280 [151,458,152,178] R Groote Eylandt 18 19 1 Northern Territory [472,976,152,178] Carpentaria -13.933333 Gulf 1289 136.6 18 19 1 Gulf R03357 R -13.966666 Groote Eylandt 1289 136.43333 Angurugu 18 19 1 Gulf R07566 [400,1066,191,217] R -13.983334 Groote Eylandt 1289 136.46666 Angurugu 18 19 1 Gulf R31879, R31881 R 22 23 1 lectotype R31880 R 22 23 1 paralectotype