Philoria knowlesi, Bertozzi, 2022

Bertozzi, Terry, 2022, A new species of Philoria (Anura: Limnodynastidae) from the uplands of the Gondwana Rainforests World Heritage Area of eastern Australia, Zootaxa 5104 (2), pp. 209-241 : 232-236

Philoria knowlesi sp. nov. Mahony MJ, Hines HB, Mahony SV, & Donnellan SC

Mt Ballow Mountain Frog

Figures 9 , 10 .

Holotype: QM J95824 an adult male collected at 820 m elevation on Ballow Creek , Mount Barney National Park, south-eastern Qld, [-28.2582, 152.6317] on 27 July 2006 by M. Mathieson and D. Ferguson.

Other material examined. See Table 1 for a full list of specimens examined.

Diagnosis. Adult size relatively small (females to 31.9 mm, males to 29.2 mm SVL); males with poorly developed nuptial pad; moderately developed facial stripe present; dorsum usually reddish brown, light tan or bronze, occasionally darker greenish brown, without larger darker blotches; conspicuous black patch on flank rarely present, a darker bar extending from across mid-thigh onto hip present in some specimens; abdomen usually pale reddish yellow, partially transparent, or darker brown with fine white spotting ( Figs 9–10 ); gular surface same as abdomen but with slightly darker hue. Ventral surfaces smooth. From a genetic perspective, apomorphic nucleotide states at five sites in the mitochondrial ND4 gene reliably diagnose P. knowlesi sp. nov. from P. loveridgei ( Table 4 ).

Dimensions of holotype (mm). SVL 25.6, HL 7.8, HW 9.4, TL 11.1, ED 2.4, EN 1.4, IN 3.1.

Description of Holotype. Body moderately flattened, robust and pear shaped. Head narrower than body, wider than long (HW/HL = 1.26), triangular in dorsal view; snout flat then rounded beyond nares, slightly sloping with a wedged tip in lateral view. Canthus rostralis moderately defined, slightly oblique; loreal region distinctly concave. Nares much nearer the eye than tip of snout, on antero-lateral edge of snout, directed laterally. Eyes moderate, diameter slightly greater than twice eye to naris distance (ED/EN = 1.66); internarial distance greater than twice the distance from eye to naris (EN/IN = 0.54). Tympana covered by skin, not conspicuous, oval shaped, vertical axis less than half eye diameter. Vomerine teeth short in two obliquely aligned plates, separated in midline, not extending laterally beyond the inner border of the choanae. Forearms and hands short, robust. Fingers short, thick, unwebbed, tips flattened dorsally, distinctly rounded ventrally, relative lengths 3>4=2>1, small subarticular tubercles beneath proximal joints, absent from distal joints, nuptial pads on first finger, comprise dense, fine conical spines, pigmented dark brown, particularly near the apex; palmar surface smooth, with prominent inner and outer metacarpal tubercles. Hindlimbs short (TL/SVL = 0.41), robust; thighs with three rows of prominent tubercles on dorsal surface. Toes unwebbed, without fringes; subarticular tubercles moderately developed; tips of toes flattened dorsally, rounded ventrally; relative lengths 4>3>5>2>1; plantar surfaces smooth, inner metatarsal tubercle flattened and rounded in shape.

Dorsal skin smooth; incomplete ridge along dorso-lateral margin, several large tubercles on upper- and midlateral surfaces; no mid-vertebral stripe; a supratympanic fold extends from behind eye over tympanum to above axilla; parotoid glands absent. Raised glandular area between posterior articulation of the jaw and anterior axil of the arm. Row of raised tubercles on posterior margin of lower jaw. Ventral surface smooth.

Variation. Adult females 26.1–31.9 mm SVL (mean = 27.9, n = 8); adult males 21.5–29.2 mm SVL (mean = 26.1 mm, n=26) ( Table 5 ). Body robust, pear-shaped. Head shorter than wide (HW/HL range 0.93–1.55), approximately one-quarter snout to vent length (HL/SVL = 0.21–0.41); snout blunt in profile, nostrils more lateral than superior. Internarial distance about double distance from eye to naris (EN/IN = 0.44–0.69). Canthus rostralis well defined, loreal region concave. Eye relatively large, diameter greater than eye to naris distance (ED/EN 1.2–2.1), pupil horizontal when constricted. Tympanum small, indistinct. Tongue approximately rectangular. Vomerine teeth short in two obliquely aligned plates, separated in midline, not extending laterally beyond the inner border of the choanae. A pale, slightly raised glandular area running posteriorly from angle of jaw o near base of forearm. Supratympanic fold distinct, often bordered by thin to moderately thick light streak dorsally, most prominent and widest on post-tympanic section of fold when present. Parotoid gland absent. An often-obscure ridge of tubercles forming lyrate pattern on dorsum originates behind eye and extends posteriorly about two-thirds along dorso-lateral margin before breaking into scattered smaller tubercules posteriorly. Dorso-lateral surfaces sometimes with 2–3 large tubercules arranged in lateral trending line; skin above eye with several tubercles. A dark patch in front of groin often occurs though usually faded to obscure, with one and sometimes two enlarged tubercles at dorsal apex of patch, not generally forming a distinct ridge or straight edge line along the dorsal side of the patch. Upper surfaces of limbs moderately tubercular.

Fingers and toes without webbing, digits short, robust, cylindrical; inner and outer palmar tubercules small but distinct ( Fig. 9C ); fingers in decreasing order of length 3>4=2>1. Hind limbs moderately long (TL/SVL = 0.35–0.46); toes in decreasing order of length 4>3>5=2>1; inner metatarsal tubercule at base of first toe small but distinct ( Fig. 9D ), no outer metatarsal tubercule. Nuptial pads on first finger, appear pale macroscopically but when examined closely comprise dense, fine conical spines, pigmented dark brown, particularly near the apex. Spatulae on first and second fingers of reproductively active females ( Fig. 10H–I ).

Colour in life. The description of variation of colour in life is based on colour photographs of eight specimens ( Fig. 10 , Table 1 ). Dorsal colour highly variable, including plain orange, orange-brown, reddish brown, brown or grey green, occasionally with suffused patches and spots of darker colour. Upper surfaces of limbs usually same colour as dorsum but a lighter hue; faint crossbars on upper thigh extend onto posterior dorsum, to produce a forward pointing v-shape marking. Inner surfaces of axils, groin and flanks, and posterior surfaces of thighs darker orangebrown than dorsum. A dark, roughly circular or irregular-shaped patch present on posterior-lateral surface, faded and obscure in some lightly coloured individuals. Toes and fingers faintly barred darker brown dorsally.

Lateral surfaces of face darker than dorsum, grading in colour intensity from the upper margin of the canthus which has a sharply demarcated black line that extends from the snout along the canthus through the eye and above the tympanum to the axil. The posterior margin behind the axil with a white margin. Upper margin of supratympanic fold lighter than dorsum; margin of upper lip spotted with white. Glandular area posterior to angle of the jaw white. Loreal region darker than dorsal colour.A black band becoming broader posteriorly from tip of snout through nostril along canthus rostralis, through eye to base of forearm. Margin along canthus rostralis forming distinct line, ventral margin is diffuse merging with the colour below. Ventral surfaces of body and hind limbs burnt orange-yellow to dark brown with pale spots, chin and throat slightly darker than abdomen without mottling. Ventral surfaces may be partially transparent so that abdominal wall and internal organs are partially visible. Iris golden dorsally and brown or black ventrally.

Etymology. Named for Mr Ross Knowles for his significant contribution to environmental conservation in Australia, and his early work on the biology and systematics of Philoria .

Distribution. Occurs in montane areas straddling the Qld-NSW border, from Levers Plateau in the east to Drummer Gully (Mount Barney National Park, Qld) in the west. All known locations are above 690 m elevation.

Habitat. In Mt Barney National Park, P. knowlesi sp. nov. occurs in simple microphyll fern forest dominated by the tree Nothofagus moorei on Cainozoic igneous rocks (Regional Ecosystem or RE 12.8.6 – Queensland Herbarium 2021), complex notophyll vine forest on Cainozoic igneous rocks (RE 12.8.5), simple notophyll vine forest with abundant Tree-ferns, Archontophoenix cunninghamiana (gully vine forest) on metamorphics +/- interbedded volcanics (RE 12.11.1). In the Lever’s Plateau area (Qld side) the rainforest community is a drier, Araucarian complex notophyll vine forest on Cainozoic igneous rocks (RE 12.8.4). Known sites in NSW are in similar rainforest habitats. The species appears to be more widespread in wetter, high-altitude rainforests where it has been recorded calling from near the summit of Mt Ballow and along the upper reaches of Lindsay Creek in NSW in Nothofagus moorei forest. In the lower drier rainforests, the species is likely confined to seepages and margins of drainages.

Biology. Very poorly known. Most specimens were collected from seepages or the boggy margins of drainage lines, habitats typical of other Philoria species. Calling has been recorded in July , and September-January inclusive. An egg mass was observed by HBH on 14 November 2006 in the headwaters of Drummer Gully , Mount Barney National Park , Qld , with an adult female in attendance. There were approximately 40 eggs in a nest chamber about 35 x 30 mm, filled with water, connected around and beneath to a porous substrate with gently seeping water, and there was no surface flow (a few eggs were lodged with the Queensland Museum - QM J84827 ). The eggs were relatively large and unpigmented and were held within a slightly frothy gelatinous mass .

Threats to habitat. A majority of the known and potentially suitable habitat for P. knowlesi sp. nov. occurs within the protected areas of Mount Barney National Park (Qld), and the NSW reserves of Mount Nothofagus National Park and the western section of Border Ranges National Park. Significant areas of Philoria habitat within Mount Barney and Mount Nothofagus National Parks burnt in the 2019 bushfires ( Heard et al. 2021 – reported as western populations of P. loveridgei ). Results of limited field surveys during the 2020/2021 and 2021/2022 breeding seasons suggest that the species has some resilience to severe drought and fire ( Heard et al. 2021), with calling males recorded at four sites within burnt catchments, including individuals calling from within habitat burnt at moderate-to-high severity. However, with greater sample sizes and survey data prior to the drought and fires, Heard et al. (2021) found reduced occupancy and abundance for P. kundagungan and P. richmondensis from nearby and similar habitat. Further, increases in fire frequency, earlier starts to fire seasons and higher intensity fires have been observed, and all are projected to proceed on the same trajectories (State of the Climate 2020; http://www. csiro.au/en /Showcase/state-of-the-climate, Dowdy et al. 2019). Although feral pigs occur in lower altitude areas of the western section of Mount Barney National Park, there is no evidence yet of them within known habitat of P. knowlesi sp. nov. However, fresh scats were found in September 2021 within 300 m of known occupied habitat (Hines pers. obs.). Feral pigs are currently severely impacting the breeding habitat of P. kundagungan in the Mistake Mountains, through wallowing and predation (Hines pers. obs.). Unauthorised cattle grazing in the western section of Mount Barney National Park also potentially threatens breeding habitat. There are few weeds of concern within known habitat at Mount Barney and Mount Nothofagus National Parks. Lantana ( Lantana camara ) is currently present at low density at some sites regenerating post-fire and has the potential to exacerbate impacts from future fires. Two significant riparian weeds are common in the region, but we have yet to observe them at sites occupied by P. knowlesi sp. nov.: Crofton weed ( Ageratina adenophora ) and mistflower ( A. riparia ). Their dense root mats bind the soil of bogs and stream banks, likely making them unsuitable for frogs to build nest chambers (Hines and Mahony pers. obs).

Figures

FIGURE 9. Philoria knowlesi sp. nov. holotype QM J95824, male from Ballow Creek, Mount Barney National Park, southeastern Queensland. A) ventral view, B) lateral view, C) snout, D) lateral view of head, E) palmar view of hand, F) plantar view of foot.

FIGURE 10. Philoria knowlesi sp. nov. in life A) QM J84821, male, Cronan Creek, Mount Barney National Park, Qld; B) QM J84820, male, Cronan Creek, Mount Barney National Park, Qld; C) QM J92852, male, Lindesay Creek, Mount Ballow, Mount Nothofagus National Park, NSW; D) QM J92851, male, Lindesay Creek, Mount Ballow, Mount Nothofagus National Park, NSW; E) AMS R185923, male, Long Creek, Levers Plateau, Border Ranges National Park, NSW; F) QM J92850, male, Lindesay Creek, Mount Ballow, Mount Nothofagus National Park, NSW; G) AMS R185923, male ventral view, Long Creek, Levers Plateau, Border Ranges National Park, NSW; H) QM J84824 female ventral view, Drummer Gully, Mount Barney National Park, Qld. Note the spatula on the second finger that is used by the female to direct bubbles of air into mucous to make a foamy egg mass; I) QM J92850 male ventral view, Lindesay Creek, Mount Ballow, Mount Nothofagus National Park, NSW; J) QM J84825 Drummer Gully, Mount Barney National Park, Qld, male ventral view.