Litoria bella Mcdonald, Rowley, Richards &amp; Frankham, 2016, sp. nov.
Creators: name: Mcdonald, Keith R.
name: Rowley, Jodi J. L.
name: Richards, Stephen J.
name: Frankham, Greta J.
Publisher: name: Zenodo
Year: 2016
Subjects: Biodiversity
Taxonomy
Animalia
Chordata
Amphibia
Anura
Hylidae
Litoria
Litoria bella

description: <i>Litoria bella</i> sp. nov. <b>Holotype</b>: QM J74476, adult male, from Lankelly Creek, 10 km NE of Coen, Cape York Peninsula, Queensland, Australia (13.8869º N; 143.2673ºE, 540 m asl; Fig. 1). Collected on 23 October 2000 by KRM. <b>Paratypes</b>: QM J74466 –8, 74470, 74473–5, 74478, eight adult males from same locality and date as holotype. QM J38289, adult male, from Little Stewart Creek, 19 km SE of Coen, Cape York Peninsula, Queensland, Australia (14.0776ºE, 143.3138º E: ~ 180 m asl), collected on 18 December 1978. QM J39656, adult male, from Coconut Creek, ~ 40 km North of Aurukun, Cape York Peninsula, Queensland, Australia (13.0082ºS, 141.7991ºE, ~ 40 m asl), collected 22 November 1981. QM J46363 –4, two adult males, Double Point Road, Shelburne Station, Cape York Peninsula, Queensland, Australia (11.9666ºS, 142.9163º E, &lt;50 m asl). <b>Etymology.</b> The specific name “ bella ” (Latin), meaning pretty or lovely, is used as an adjective in reference to the beautiful coloration of this species. <b>Suggested common name.</b> Cape York Graceful Treefrog. <b>Diagnosis.</b> The new species is assigned to the genus <i>Litoria</i> based on the presence of a horizontally oriented contracted pupil and the absence of a palpebral reticulum (Zweifel 1958; Tyler &amp; Davies 1978). Based upon molecular and morphological data the new species is closely related to <i>Litoria auae</i>, and it exhibits other characters typical of the <i>L. gracilenta</i> group: moderately small size (&lt;50 mm SVL), green dorsum and a male advertisement call resembling a low-pitched growl (Menzies 2006; Menzies &amp; Tyler 2004). However it is distinguished from all congeners by the following combination of characters: size moderate (male body size 34.5–41.8 mm), dorsum near-immaculate green, venter orange, digits and webbing bright orange, lateral surfaces of the thigh bluish purple, no pale canthal stripe, bones white, and male advertisement call a finely pulsed single note with a pulse rate of 56– 64 pulses/s and dominant frequency of 2.6–2.8 kHz at 21.4ºC. <b>Description of holotype.</b> Adult male, moderate body size (38.5 mm SVL; Fig. 3); head length equal to head width (101%); snout obtusely rounded in dorsal view and truncate in profile; canthus rostralis poorly defined; loreal region flat; interorbital region flat; nostrils oval, closer to tip of snout than eye; eyes moderately large (10% SVL), pupil oval, horizontal, tympanic rim slightly elevated relative to skin of temporal region, top margin obscured by supratympanic skin fold, tympanum 64% of eye diameter; vomeropalatines with two patches of teeth (one on each side) between internal nares (Fig. 3). Vocal sac present; slightly raised supratympanic fold extending to just beyond level of axilla. Finger webbing does not extend to disks (Fig. 3 C), relative length of fingers I &lt;II &lt;IV &lt;III; tips of all fingers with well-developed disks with distinct circummarginal grooves, disks relatively wide compared to penultimate phalanx width (third finger disk 143% third finger width); subarticular tubercles indistinct; nuptial pads present on posteriodorsal surface of finger I. Relative length of toes I &lt;II &lt;V &lt;III &lt;IV; tips of toes with well-developed disks with distinct circummarginal grooves; disks slightly smaller than those of fingers (fourth toe disk 131% fourth penultimate phalanx width); toe webbing on fourth and fifth toe reaches base of disk, webbing between first and second toe reaches edge of terminal subarticular tubercle (Fig. 3 D); subarticular tubercles indistinct, rounded; inner metatarsal tubercle low, oval; outer metatarsal tubercle absent. Tibia length moderately long (55% SVL). Skin of dorsal surfaces finely granular; ventral surfaces of body and thighs coarsely granular, finer on chest and under arms, smooth under shanks. <b>Colour of holotype in preservative.</b> Dorsal ground colour uniform greyish green. Venter, upper arms, inner surface of lower arms, hands and feet including webbing pale yellowish orange. Upper thighs pale yellow with purplish brown wash. Iris pale yellow with black periphery. <b>Measurements (mm).</b> Holotype: SVL 38.5, TIB 21.2, HL 13.8, HW 13.7, EYE 3.7, TMP 2.4, TEY 1.2, END 3.6, IND 3.4, 3FD 2.0, 4TD 1.7. <b>Colour in life.</b> Recorded from images of the species from the Iron and McIlwraith Ranges (Fig. 4). Dorsal surface bright green, with bright orange on the dorsal surface of the upper arms, fingers I–III, toes I–IV and distal end of finger IV and toe V, including webbing and toe-disks. Blueish purple inner and outer thighs with only thin strip of green along centre. Pale canthal stripe absent; paler yellowish along bottom of upper lip and along flanks. Ventral surfaces are orange, more yellowish along outer edges of limbs and belly. Iris bright yellowish gold with a network of fine darker gold reticulations; iris periphery black. <b>Variation.</b> The type series exhibits little variation in colour or morphology, and specimens observed in life consistently exhibited the colours described above. The large variation in extent of dorsal spotting that is apparent in many other members of the <i>L. gracilenta</i> group is not apparent in <i>L. bella</i>. Adult males vary in size from ~ 34–41 mm but variation in morphological features is otherwise slight: measurements of the type series are shown in Table 4. continued. <b>Advertisement call.</b> Call descriptions are based on the calls of two topoparatypes (QM J74467 and QM J74468), recorded at an air temperature of 24.1ºC within 0.3 m of calling sites (Table 5; Fig. 5). The advertisement call of <i>Litoria bella</i> consists of a single note, although three of the eight recorded calls of QMJ 74467 were only separated by a short interval. The duration of each call varied from 1.42– 2.62 s and calls were separated by an interval of 0.88– 5.78 s. Each call had 89–156 uniformly spaced pulses repeated at a rate of around 60 pulses/s. Calls were amplitude modulated, increasing gradually to a peak at approximately 70% of the call duration (except for the three calls separated by a short interval). The dominant frequency was 2.6–2.8 Hz, but frequency increased over the duration of the call about 0.2 kHz, from 2.5–2.6 to 2.7–2.8 kHz. <b>Ecology.</b> <i>Litoria bella</i> inhabits moist monsoon rainforest and vine thicket on hills and plains. The species has been observed breeding in ephemeral pools adjacent to permanent streams and observed (but not breeding) around ephemeral pools in gullies flowing into permanent streams. In October 2000, breeding was observed after a thunderstorm. Two pairs of frogs in inguinal amplexus (female QM J74465 and male QM J4466, and QM J74469 and QM J74470) laid spawn with 606 and 844 dorsally brown-pigmented eggs in clear jelly respectively. <b>Distribution and conservation status.</b> <i>Litoria bella</i> is known from 5–560 metres elevation between Moa Island in the north and about 20 km south of Coen in the south (Fig. 1). The species appears to be patchily distributed within the Cape York Peninsula Bioregion (Sattler &amp; Williams 1999), while <i>Litoria gracilenta</i> is restricted to the Wet Tropics, Brigalow Belt, Central Queensland Coast and South East Queensland Bioregions and the Eastern Australian Temperate Forests (Sattler &amp; Williams 1999). The southern-most location of the new species is separated by 220 km from the most northern record of <i>Litoria gracilenta</i> in the Endeavour Valley. <i>Litoria bella</i> is known to occur within at least three protected areas, the Kulla National Park (CYPAL), Kutini-Payamu (Iron Range) National Park (CYPAL) and the Oyala Thumotang National Park (CYPAL). Given its relatively broad distribution, no known threats, and its occurrence within at least three protected areas the species likely falls under the IUCN classification of Least Concern. <i>Litoria bella</i> <b>sp. nov.</b> <i>Litoria gracilenta</i> QM J74467 QM J74468 AMS R184768 AMS R184777 Measured 8 calls (3 double=5 sets) 6 calls (all single) 6 calls 6 calls Call length (s) 1.94 (1.42–2.62) 2.34 (2.29–2.43) 2.06 (2.01–2.09) 2.20 (1.31–2.64) Intercall interval (s) 2.85 (0.88–4.54) 5.78 (5.05–5.78) 3.38 (2.62–4.49) 5.39 (4.18–7.33) Pulses 116.5(89–156) 140.8 (135–149) 180.2 (176–191) 183 (116–225) Pulse/s 60.6(55.8–64.0) 59.9(57.7–63.6) 85.7(85.7–93.0) 84.2(74.7–88.8) Dominant Frequency (Hz) 2.7 (2.7–2.8) 2.6 (2.6–2.7) 2.4 2.7 (2.6–2.8) Temperature (ºC) 24.1 24.1 21.8 21.7 <b>Comparisons.</b> From the eight species of the <i>Litoria gracilenta</i> group currently known, the new species may be distinguished by the combination of (1) male body size moderately large (34.5–41.8 mm SVL), (2) nearimmaculate green dorsum, (3) orange venter, (4) bright orange digits and webbing, (5), iridescent bluish purple lateral surfaces of thigh, (6) no pale canthal stripe, (7) white bones, and (8) a highly-pulsed, single-note, male advertisement call with a pulse rate of 56–64 pulses/s and dominant frequency of 2.6–2.8 kHz at 21.4ºC. From <i>L. auruensis</i>, the new species further differs by lacking broad lateral fringes on fingers and toes (vs present in <i>L. aruensis</i>), and in having white bones (versus green in <i>L. aruensis</i>). It further differs from <i>L. auae</i> (Fig. 6 A) by having a larger (although slightly overlapping) male body size (34.5–41.8 mm SVL vs 29.9–36.2 mm in 37 <i>L.</i> auae), smaller relative eye size (0.10–0.12 EYE/SVL vs 0.12–0.16 in <i>L. auae</i>), bright orange webbing (vs golden yellow in <i>L. auae</i>), immaculate dorsum in most individuals, with only a few small spots on the dorsum on five of 13 types (vs few to many small, scattered pale yellow spots in all but a few of the 40 types of <i>L. auae</i>), bluish purple dorsal thigh coloration (vs yellow in <i>L. auae</i>), no pale canthal stripe (vs indistinct yellow canthal stripe in <i>L. auae</i>), relatively longer (although slightly overlapping) male advertisement call (1.42– 2.62 s at 24.1ºC vs 0.48– 1.78 s at 24ºC in <i>L. auae</i>), and molecular divergence (see below). The new species differs from <i>L. callista</i> by having a larger male body size (34.5–41.8 mm SVL vs 31.5–34.5 mm in <i>L. callista</i>), orange venter (vs yellow <i>in L. callista</i>), bluish purple dorsal thigh coloration (vs orange in <i>L. callista</i>), no pale canthal stripe (vs pale canthal stripe in <i>L. callista</i>), bright orange webbing (vs yellow in <i>L. callista</i>), and a male advertisement call of a longer duration (1.42– 2.62 s at 24.1ºC vs 0.045– 0.190 s at 24ºC in <i>L. callista</i>), greater average number of notes per call (~117–141 vs 9.5 in <i>L. callista</i>) and a higher dominant frequency (2.6–2.8 kHz at 24.1ºC vs 1.28–2.39 kHz at 24ºC in <i>L. callista</i>). From <i>L. elkeae</i>, the new species differs by having a larger male body size (34.5–41.8 mm SVL vs 27.5–30.4 mm in <i>L. elkeae</i>), immaculately green dorsum in most individuals (vs pale green with indistinct whitish dorsal spotting in <i>L. elkeae</i>), orange venter (vs cream in <i>L. elkeae</i>), no pale canthal stripe (vs white canthal stripe in <i>L. elkeae</i>), and a male advertisement call with a single note with a relatively slow pulse rate (56–64 pulses/s at 24.1ºC vs a biphasic call with 80–101 pulses/s in <i>L. elkeae</i>). From <i>L. eschata</i>, the new species differs by having an orange venter (vs yellow in <i>L. eschata</i>), bluish purple dorsal thigh coloration (vs caramel brown in <i>L. eschata</i>), a male advertisement call with a slower pulse rate (56–64 pulses/s at 21.4ºC vs 72–80 in <i>L. eschata</i>) and higher dominant frequency (2.6–2.8 kHz at 24.1ºC vs 2.2–2.5 kHz in <i>L. eschata</i>). From <i>L. gracilenta</i> (Fig. 6 B–D), the new species differs by having an orange venter (vs white to yellow in <i>L. gracilenta</i>), bluish purple dorsal thigh coloration (versus purplish red in <i>L. gracilenta</i>), no pale canthal stripe (vs pale yellowish canthal stripe in <i>L. gracilenta</i>), orange webbing (vs yellow in <i>L. gracilenta</i>), bright golden iris with thick black iris periphery (vs brownish gold iris, often with blue or green at borders and a narrow black periphery in <i>L. gracilenta</i>), a male advertisement call with fewer pulses (~120–140 at 24.1ºC vs ~180 pulses in <i>L. gracilenta</i> at 21.7–21.8ºC) and a slower pulse rate (~60 at 24.1ºC vs ~85 pulses/s in <i>L. gracilenta</i> at 21.7–21.8ºC), and molecular divergence (see below). From <i>L. kumae</i>, the new species differs by having a larger male body size (34.5–41.8 mm SVL vs 24–30 mm in <i>L. kumae</i>), an orange venter (vs a venter that is white anteriorally and yellowish gold posteriorally in <i>L. kumae</i>), bluish purple dorsal thigh coloration (vs reddish gold in <i>L. kumae</i>) and no pale canthal stripe (vs pale canthal stripe in <i>L. kumae</i>). From <i>L. robinsonae</i>, the new species differs by having a larger male body size (34.5– 41.8 mm SVL vs 28.3–28.7 mm in <i>L. robinsonae</i>), orange webbing and digits (vs bluish-grey webbing with bluishgrey to translucent anterior digits) and no pale canthal stripe (vs pale canthal stripe in <i>L. robinsonae</i>). The new species differs from <i>Litoria chloris</i> (Fig. 6 E) by having a smaller male body size (34.5–41.8 mm SVL vs&gt; 45 mm), an orange venter (versus yellowish white in <i>L. chloris</i>), orange webbing (vs yellow in <i>L. chloris</i>), bright golden iris with thick black iris periphery (vs reddish-orange with thick black periphery in <i>L. chloris</i>), a male advertisement call of a single, pulsed note of more than 1s duration (vs a multi-note call with each note &lt;1s in duration in <i>L. chloris</i>; see Fig 5 C), and molecular divergence (see below). The new species differs from <i>Litoria xanthomera</i> (Fig. 6 F) by having a smaller male body size (34.5–41.8 mm SVL vs&gt; 45 mm), bright orange webbing (vs yellow to pale orange-yellow in <i>L. xanthomera</i>), bright golden iris with thick black iris periphery (vs reddishorange with thick black periphery in <i>L. xathomera</i>), bluish purple dorsal thigh coloration (vs orange in <i>L. xanthomera</i>), and a male advertisement call of a single, pulsed note of more than 1s duration (vs a multi-note call with each note &lt;1s in duration in <i>L. xanthomera</i>). Tyler and Davies (1978) included the New Guinea species <i>Litoria graminea</i> and <i>L. multiplica</i> in their <i>L. aruensis</i> group. <i>Litoria graminea</i> and the closely related <i>L. dux</i> and <i>L. sauroni</i> are large (male SVL&gt; 60 mm), robust canopy dwelling frogs with prominent white dermal ridges on the arms and legs and are restricted to New Guinea (Richards &amp; Oliver 2006). <i>Litoria multiplica</i> is a similar size to <i>L. bella</i> but has a prominent white bilobed dermal ridge around the vent, white dermal ridges around the knees and along the outer edge of the tarsus, and dark purplish black lateral spots or mottling (Richards <i>et al.</i> 2009). <b>Sequence divergence.</b> Uncorrected sequence divergences between <i>L. bella</i> and the two most morphologically similar species, <i>L. gracilenta</i> and <i>L. auae</i>, was 20.1–23.0% and 13.0–14.5% at the more rapidly evolving ND4 gene fragment analysed and 5.2–6.1% and 2.0% at the relatively conservative 16S gene fragment analysed. Uncorrected sequence divergences between <i>L. bella</i> and <i>L. chloris</i> was 20.5–21.5 % (ND4) and 4.8–5.0% (16S). These values are comparable to those reported among other recognised <i>Litoria</i> species (Mahony <i>et al.</i> 2001; Burns &amp; Crayn 2006; Smith <i>et al</i>. 2013).
description: Published as part of <i>Mcdonald, Keith R., Rowley, Jodi J. L., Richards, Stephen J. &amp; Frankham, Greta J., 2016, A new species of treefrog (Litoria) from Cape York Peninsula, Australia, pp. 153-169 in Zootaxa 4171 (1)</i> on pages 159-166, DOI: 10.11646/zootaxa.4171.1.6, http://zenodo.org/record/268260
description: {"references": ["Tyler, M. J. &amp; Davies, M. (1978) Species-groups within the Australopapuan hulid frog genus Litoria Tschudi. Australian Journal of Zoology, 26, 1 - 47. http: // dx. doi. org / 10.1071 / AJZS 063", "Menzies, J. (2006) The frogs of New Guinea and the Solomon Islands. Pensoft Publishers, 345 pp.", "Menzies, J. I. &amp; Tyler, M. J. (2004) Litoria gracilenta (Anura: Hylidae) and related species in New Guinea. Australian Journal of Zoology, 52, 191 - 214. http: // dx. doi. org / 10.1071 / ZO 03008", "Sattler, P. S. &amp; Williams, R. D. (1999) The Conservation Status of Queensland Bioregional Ecosystems. Environmental Protection Agency, Brisbane.", "Richards, S. J. &amp; Oliver, P. (2006) Two new species of large green canopy-dwelling frogs (Anura: Hylidae: Litoria) from Papua New Guinea. Zootaxa, 1295, 41 - 60.", "Richards, S. J., Oliver, P., Krey, K. &amp; Tjaturadi, B. (2009) A new species of Litoria (Amphibia: Anura: Hylidae) from the foothills of the Foja Mountains, Papua Province, Indonesia. Zootaxa, 2277, 1 - 13.", "Mahony, M., Knowles, R., Foster, R. &amp; Donnellan, S. (2001) Systematics of the Litoria citropa (Anura: Hylidae) complex in northern New South Wales and southern Queensland, Australia, with the description of a new species. Records of the Australian Museum, 53, 37 - 48. http: // dx. doi. org / 10.3853 / j. 0067 - 1975.53.2001.1322", "Burns, E. L. &amp; Crayn, D. M. (2006) Phylogenetics and evolution of bell frogs (Litoria aurea species-group, Anura: Hylidae) based on mitochondrial ND 4 sequences. Molecular Phylogenetics and Evolution, 39, 573 - 579. http: // dx. doi. org / 10.1016 / j. ympev. 2005.11.017", "Smith, K. L., Hale, J. M., Kearney, M. R., Austin, J. J. &amp; Melville, J. (2013) Molecular patterns of introgression in a classic hybrid zone between the Australian tree frogs, Litoria ewingii and L. paraewingi: evidence of a tension zone. Molecular Ecology, 22, 1869 - 1883. http: // dx. doi. org / 10.1111 / mec. 12176"]}