Molecular systematic analysis demonstrates that the threatened southern bell frog, Litoria raniformis (Anura: Pelodryadidae) of eastern Australia comprises two sub-species Author Vörös, Judit Department of Zoology, Hungarian Natural History Museum H- 1088 Budapest, Baross u. 13, Hungary Author Wassens, Skye 0000-0001-8886-8426 School of Agriculture, Environment and Veterinary Sciences, Charles Sturt University, PO Box 789, Albury, 2640, Australia. swassens @ csu. edu. au; https: // orcid. org / 0000 - 0001 - 8886 - 8426 swassens@csu.edu.au Author Price, Luke School of Biological Sciences, University of Adelaide, North Terrace, Adelaide 5005, Australia. Author Hunter, David New South Wales Department of Planning, Industry and Environment, Albury, 2640, Australia. David. hunter @ environment. nsw. gov. au Author Myers, Steven 0000-0001-8885-8770 South Australian Museum, North Terrace, Adelaide, 5001, Australia. & Steven. Myers @ alsglobal. com; https: // orcid. org / 0000 - 0001 - 8885 - 8770 yers@alsglobal.com Author Armstrong, Kyle School of Biological Sciences, University of Adelaide, North Terrace, Adelaide 5005, Australia. & South Australian Museum, North Terrace, Adelaide, 5001, Australia. Author Mahony, Michael J. 0000-0002-1042-0848 School of Environmental and Life Sciences, University of Newcastle, University Drive, Callaghan NSW 2308, Australia michael. mahony @ newcastle. edu. au; https: // orcid. org / 0000 - 0002 - 1042 - 0848 michael.mahony@newcastle.edu.au Author Donnellan, Stephen 0000-0002-5448-3226 South Australian Museum, North Terrace, Adelaide, 5001, Australia. & Steve. donnellan @ samuseum. sa. gov. au; https: // orcid. org / 0000 - 0002 - 5448 - 3226 teve.donnellan@samuseum.sa.gov.au text Zootaxa 2023 2023-01-11 5228 1 1 43 journal article 222253 10.11646/zootaxa.5228.1.1 7b9a994e-a6ec-4efb-a755-e1e8789a2ee0 1175-5326 7524042 DA207B1D-81A2-4176-AA9A-64B1D484C307 Litoria raniformis major ( Copland, 1957 ) Figs 14 , 15 Neotype . TMAG C290 , an adult female from Lauderdale , Tasmania , Australia . 42.91° S , 147.49° E . Collected by Mr Van Der Water on 27April 1971 . Diagnosis. Litoria raniformis major can be diagnosed from L. r. raniformis and from L. castanea by the presence of 35 and 34 diagnostic nucleotide sites in the ND4 alignment respectively ( Table 3 ). Measurements of neotype (mm). SVL: 65.8, HL: 23.9, HW: 24.7, IND: 4.2, NS: 5.4, IOD: 4.4, EN: 6.21, ED: 5.26, TD: 5.3, FeL: 33.9, TL: 32.5, FoL: 47.4, FLL: 12.6, RLF: 29.6, D1T: 1.7, D4T: 1.9, D1F: 1.7, D3F: 1.8. Description of Neotype . Snout prominent, rounded when viewed from above and in profile. Nostrils more lateral than superior, closer to snout than to eye. Canthus rostralis well defined and straight. Eye relatively large (ED/HL 0.22). Tympanum distinct, circular, length about equal to eye diameter (TD/ED 1.2). Vomerine teeth short straight plates bridging the gap between the choanae. Tongue approximately rectangular. Fingers long, broad, webbing absent. Subarticular tubercles prominent, palmar tubercles not prominent. Terminal discs not prominent, barely extending beyond lateral extremities of penultimate phalanx. Fingers in order of length 3>4>1>2. Hindlimb length moderate (TL/SVL 0.49). Toes in order of length 4>5=3>2>1. Webbing on toes I, II, III, V reaches base of terminal toe disc, and to base of the penultimate phalange on toe IV ( Fig. 14D ). Subarticular tubercles not prominent. Oval inner metatarsal tubercle prominent, approximately one-quarter length of first toe. Terminal toe discs not prominent, not extending beyond lateral extremities of penultimate phalanx. Variation. A summary of variation in 17 mensural traits is presented in Table 6 . Mean SVL: females = 71.5 mm , males = 62 mm . Head length equal to head width (HL/HW 0.92–1.17) and approximately one-third of SVL (HL/SVL 0.3–0.39). Distance between eye and naris equal to or greater than internarial span (EN/IND 1.03–1.45). Eye relatively large, its diameter variable relative to eye to naris distance (ED/EN 0.85-1.54). Pupil horizontal when constricted ( Fig. 15 ). Tympanum length greater than half eye diameter (TD/ED 0.63–1.17). FIGURE 14 . Neotype designated for Litoria r. major ( Copland, 1957 ) TMAG C 290. A ) dorsal, B ) ventral and C ) lateral view, D ) plantar view of foot, E ) palmar view of hand. Hindlimb length moderate (TL/SVL 0.42–0.54). Toe subarticular tubercles not prominent. Oval inner metatarsal tubercle prominent, approximately one-fifth to one-sixth length of first toe. Dorsum mildly granular with varying coverage of low tubercles. Upper surface of limbs smooth or varying coverage of low tubercles. Flanks with dense coverage of low tubercles. Abdomen, undersurface of thighs, and lateral aspect of body coarsely granular. Distinct pectoral fold present. Small vocal slits present in buccal cavity at base of tongue parallel to lateral margin of tongue. Colour pattern. Variation in colour is described from images taken in life ( Fig. 15 ). Head smooth with uniform colour. Iris gold. Canthus rostralis typically uniformly colored with the light tones of the dorsum or with patches of light green. Darker tympanum contrasts with lighter loreal region. Green to bronze mid-dorsal stripe extends from the snout or from between the eyes to the cloaca varying in width and prominence. Back covered in prominent tubercles arranged either in lines or irregularly that are in some cases enhanced with dark colouration. Dorsal colour varies from green to bronze sometimes with darker patches. A clearly demarked line with a black lower margin runs from the nostril through the eye, above the tympanum and then along the dorso-lateral margin to the groin. In the dorso-lateral region its upper margin is enhanced by prominent cream or white or green tubercles. Flanks with prominent cream or white coloured coarse granules or tubercles. Colour pattern of limbs is a continuation of dorsal colour and pattern, leading edge of lower leg usually with dark margin. Back of thighs awash with emerald green to blue, moderately granular proximally with few to moderate numbers of prominent light coloured tubercles. Belly and chin cream or white, undersurfaces of limbs either cream or with light green flush, with dark patches. Groin regions with similar pattern and colour to back of thigh. Throat dusky in mature males. FIGURE 15 . Images in life of Litoria r. major . A ) Werribee, Vic, Stephen Mahony; B ) Werribee, Vic, Stephen Mahony; C) Neerim East, Gippsland, Vic, Peter Robertson; D) Darebin Creek, Thomastown, Vic., Peter Robertson; E ) Montana, Tasmania, Ryan Francis; F ) Westbury, Tasmania, Ryan Francis. Distribution. Southern Victoria , south-eastern New South Wales , south-eastern South Australia and Tasmania . It occurs in the IBRA regions: Murray Darling Depression, South East Corner, South East Coastal Plain, Southern Volcanic Plain, Victorian Midlands, Naracoorte Coastal Plain, Furneaux, King, Ben Lomond, Tasmanian Central Highlands,Tasmanian Northern Midlands, Tasmanian Northern Slopes, and Tasmanian South East.Our mitochondrial genetic data identified a L. r. major (SAMA R15375 A) at a location, 4 miles S Verdun, in the central Mount Lofty Ranges south-east of Adelaide (collection date not available). A genetically identified L. r. raniformis was also collected at this locality (SAMA R15375 B). TABLE 6. Summary of variation in 19 mensural traits in the northern and southern groups of L. raniformis . Mean±SD and ranges are presented. ED and EN were measured from fewer vouchers as follows: southern group 5 males, 8 females; northern group 6 males, 7 females.

sex	females	males
taxon	southern	northern	southern	northern
r. major	r. raniformis	r. major	r. raniformis
N	38	31	62	18
SVL	71.46±10.06	69.26±8.25	60.46±6.73	61.74±6.12
55.1–94.2	46–78.8	44.8–74.5	48.7–69.7
HL	24.55±3.24	24.34±2.69	21.18±2.17	21.48±1.64
19.5–35	17.4–28.2	16.9–26.1	18.1–23.7
HW	24.51±3.43	25.2±2.77	21.2±2.23	22.3±2.3
19.4–34.9	18.2–30.3	17.3–27.1	17.2–25.2
SL	12.99±1.7	12.78±1.4	11.27±1.31	11.65±0.82
10.5–18.2	9.5–14.9	8.6–15.6	9.8–13
IND	3.89±0.53	3.77±0.51	3.4±0.41	3.46±0.36
2.9–5	2.9–4.9	2.7–4.8	2.5–3.9
NS	5.04±0.77	4.91±0.75	4.32±0.6	4.25±0.43
3.8–7.6	3–6.2	3.1–5.7	3.7–5.1
IOD	4.02±0.69	4.31±0.64	3.39±0.5	3.61±0.46
2.6–6	2.8–5.9	2.4–5.5	3–4.4
EN	5.44±0.68	5.73±0.5	4.72±0.32	5.14±0.35
4.41–6.34	5.17–6.66	4.44–5.25	4.59–5.55
ED	6.67±1.23	6.56±0.5	5.97±0.67	6.44±0.5
4.54–8.13	5.87–7.12	5.11–6.84	5.93–7.05
TD	5.33±0.79	5.19±0.58	4.57±0.49	4.79±0.42
4–7.5	3.7–6.1	3.8–5.9	4.1–5.7
FeL	33.3±4.37	32.4±4.23	28.43±3.64	29.21±3.95
26.8–44.6	21.2–40.9	21.3–38.7	22.2–35.9
TL	33.68±3.52	33.08±3.93	29±3.02	30.25±2.96
26.4–39.7	22.6–39.7	22–37	25.6–35.9
FoL	48.57±5.88	46.38±6.16	42.17±4.87	42.33±4.84
38.6–58.2	30.4–57.5	31.7–52.2	32.8–48.5
FLL	12.29±1.72	11.21±1.55	10.8±1.8	10.57±1.1
8.5–15.8	7.5–13.5	7.1–16.1	8.7–12.7
RLF	29.9±3.75	29±3.58	25.54±2.56	26.34±3.07
23.8–37.4	19.7–35.6	18.9–31.5	20.4–30.4
D1T	1.5±0.26	1.45±0.29	1.32±0.2	1.28±0.24
0.8–2.1	0.8–2.1	1–1.8	0.9–1.6
D4T	1.75±0.33	1.58±0.31	1.5±0.24	1.49±0.19
0.9–2.4	1–2.5	0.9–2.1	1–1.7
D1F	1.43±0.27	1.55±0.47	1.29±0.22	1.31±0.16
1–2.2	1–3.5	0.8–1.8	1.1–1.6
D3F	1.62±0.27	1.55±0.3	1.47±0.23	1.38±0.19
0.9–2.3	0.9–2.1	1–2.1	1.1–1.8

Ecology and reproductive biology. Litoria r. major occurs in cooler mesic habitats. Breeding occurs in spring and summer, in permanent and ephemeral wetland. Preference is shown for wetland habitats with a large proportion of emergent, submerged and floating vegetation, and slow-flowing or still water ( Robertson et al . 2002 , Heard et al. 2004 , 2008 , Hamer & Organ 2008 , Clemann & Gillespie 2012 ). Demographic studies indicate a metapopulation structure around separate or interconnected wetlands, with permanent waterbodies, or those near permanent water, favoured ( Heard et al . 2010 , Clemann & Gillespie 2012 ). In disturbed areas it occupies artificial waterbodies such as farm dams, irrigation channels, and disused quarries. Frogs overwinter beneath thick vegetation, logs, rocks and other ground debris, sometimes at considerable distances from waterbodies. Breeding does not appear to be as reliant upon flooding as in L. r. raniformis ( White & Pyke 1999 ) but see Robertson et al. (2002) and Heard et al . (2004) . Breeding occurs in spring and summer, and the larval phase is often longer than in L. r. raniformis ( Turner 2022 ) . Anstis (2017) described the eggs and larval stages and development of L. r. major from south-eastern Victoria and Tasmania.