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