Philoria loveridgei Parker, 1940

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 : 229-230

Philoria loveridgei Parker, 1940

Figures 7 , 8 .

Holotype. BMNH 1933.4.8.6, by original designation; renumbered BMNH 1947.2.1994 according to Cogger et al. (1983). Adult female.

Type locality. " McPherson Range, 3–4,000 ft, South Queensland ", Australia .

Shea (2005) discussed the itinerary of J. P. Darlington, the collector, and suggested that the type locality lies within Lamington National Park and was accessed from O’Reilly’s Guest House that was opened in 1926. The collection location was most likely along the “Border Track” which runs south-east from the guest house and passes through cool upland rainforests at an altitude of ~ 920–1200m asl (= 3018–3937 ft), where Philoria are currently known from several locations .

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

Diagnosis. Adult size relatively small (SVL to 33 mm); males with poorly developed nuptial pad on first finger; black head stripe well-developed; flanks either entirely black or with a black mark of variable size; and dorsum either brown, reddish-brown bronze or light grey. From a genetic perspective, apomorphic nucleotide states at five sites in the mitochondrial ND4 gene reliably diagnose P. loveridgei from P. knowlesi sp. nov. ( Table 4 ).

Dimensions of holotype (mm). SVL 29.9, HL 7.2, HW 10.5, TL 12.9, ED 3.0, EN 1.3, IN 3.4.

Description of the Holotype. Body moderately flattened, robust and pear shaped. Head narrower than body, wider than long (HW/HL = 1.45), triangular in dorsal view; snout flat then rounded beyond nares, slightly sloping with wedged tip in lateral view, blunt in profile; canthus rostralis well defined, slightly oblique; loreal region distinctly concave. Nostrils more lateral than superior, much nearer eye than tip of snout, on antero-lateral edge of snout, with raised rim on anterior margin. Eyes moderately large, diameter slightly greater than twice eye to naris distance (ED/EN = 2.33); internarial distance greater than twice the distance from eye to naris (EN/IN = 0.57). Tympanum covered by skin, not conspicuous, oval shaped. Vomerine teeth short in two obliquely aligned plates, separated in midline, not extending laterally beyond the inner border of the choanae. Tongue broad and rectangular. Forearms and hands short and robust. Fingers short and thick, unwebbed, tips flattened dorsally, distinctly rounded ventrally, relative lengths 3>4>2>1, subarticular tubercles beneath the proximal joints small. Palmar surface smooth, with a poorly defined elongate flattened inner metatarsal tubercle. Hindlimbs short (TL/SVL = 0.43), robust. Toes unwebbed, without fringes; subarticular tubercles poorly developed, relative lengths 4>3>5>2>1; tips of toes flattened dorsally, rounded ventrally, some tips sunken and spatulate due to preservation; plantar surfaces smooth; small inner, but no outer metatarsal tubercule. Dorsal skin smooth, without parotoid gland or ridges; a supratympanic fold from behind eye to above axilla; ventral skin smooth; mid-vertebral stripe absent.

Variation. Two adult females 28.9 and 29.7 mm SVL (mean 29.3 mm) and 53 adult males 22.8–33.0 mm (mean 27.02 mm) ( Table 5 ). Head shorter than wide (HW/HL = 1.09–1.74), approximately one quarter snout-vent length (HL/SVL range = 0.20–0.31). Internarial distance about twice the distance from eye to naris (EN/IN = 0.43–0.69). Eye relatively large, its diameter greater than eye to naris distance (ED/EN = 0.91–2.3). Pupil horizontal when constricted.

A slightly raised glandular area from corner of mouth posterior to near base of forearm evident in all specimens, often a pale colouration to that of the loreal surface ( Fig. 8A–F ). Supratympanic fold distinct, often bordered by thin pale streak dorsally, especially on post-tympanic section. Dorsal surfaces smooth or finely tubercular ( Fig. 8G & H ). A pair of raised ridges starting at posterior corner of eye and extending posteriorly half-way to two-thirds along dorsum obvious in most specimens but sometimes obscure ( Fig. 8A–C ). Posteriorly this ridge sometimes continues as disjunct large tubercules ending above groin and forming an overall lyrate pattern. Additional scattered tubercules may also be present dorso-laterally, mid-dorsally and on skin above eye. Upper surfaces of limbs moderately tubercular.

Hind limbs short (TL/SVL = 0.38–0.48). Nuptial pad on upper surface of first finger of adult male consists of dense, fine conical spines, pigmented dark brown, more intensely near their apex. Spatulae present on first and second fingers of reproductively active females ( Fig. 8J ).

Colour in life. The description of variation of colour in life is based on colour photographs of seven specimens ( Fig. 8 ). Dorsal colour varies from light grey mid-dorsal area with black flanks and upper surfaces of limbs, to light brown with darker brown irregular patches, to fawn with a few spots of darker pigment and a black or dark brown patch on flank, to uniform dull claret on upper surfaces and flanks ( Fig. 8 ). In many specimens’ posterior third of dorsal surface above hips and on upper surface of legs more darkly pigmented than anterior dorsal surface. Limbs sometimes with faint transverse bands. A dark brown or black stripe extends from snout, through eye, to base of arm. A dark patch usually occurs on the dorsal surface of the thigh which is continuous with a patch on posterior dorsal surface of the body when the legs are adpressed, usually obvious on lightly coloured individuals, but can be indiscernible on very dark individuals. The posterior edge of the patch is generally straight in most individuals and in line with a tuberculate ridge, with the colouration fading posterior to the tubercule. Colour of ventral surfaces of body, limbs and throat highly variable ( Fig. 8G–I ), from uniformly pale to darkly pigmented with numerous white or cream spots. The abdomen ranges from transparent or with pale or orange-brown pigmentation. A pair of small dark marks either side of vent often present.

Distribution. Philoria loveridgei occurs on the Tweed caldera, a landform comprising the eroded caldera and central cone of the ancient Tweed Volcano. The northern rim of the caldera is the eastern McPherson Range that includes the Lamington and Springbrook Plateaus in Qld, while the western rim forms the Tweed Range in NSW, and the southern rim the Nightcap Range in NSW, with Wollumbin in NSW the central cone of the volcano. The Tweed shield volcano was active about 23 million years ago for a period of about 3 million years ( Knesel et al. 2008). Subsequently, the caldera was incised deeply by streams that flowed radially, resulting in the breaks between the named ranges and plateaus, while the basin of the caldera and the eastern caldera rim were eroded by the Tweed River with tributaries that originate on, and partially surround Wollumbin. Our genetic analysis indicates that the populations on each of these mountain ranges can be distinguished as distinct lineages ( Figs 3–5 ), but not to a level that warrants the recognition of distinct species. The distinction among the lineages suggests that gene flow is limited due to low dispersal between the upland plateaus and across the river valleys. It is also likely that climate fluctuations in the past resulted in the expansion and contraction of rainforest communities in the valleys and on the high mountains of the region where most populations of Philoria are found ( Bryant & Krosch 2016).

Figures

FIGURE 3. Phylogenetic analyses of SNP data from northern Philoria. A) Maximum likelihood IQTree with bootstrap proportions from 100 pseudo-replicates from a fast search. B) SVD Quartets tree with bootstrap proportions from 100 pseudo-replicates with full search.

FIGURE 4. Bayesian mitochondrial ND4 phylogram with maximum likelihood bootstrap proportions (left) and Bayesian posterior probabilities (right) values at each node. The tree was rooted with sequences for the outgroup P. frosti but we excluded these so that the tree could be presented at a readable scale due to the long branch to the outgroup. Numbers at start of each terminal taxon name refer to locality code in Table 1.

FIGURE 5. Population structure analysis of the SNP data for A) P. kundagungan and B) P. loveridgei. For collection locality site numbering refer to Table 1 and Fig. 1.

FIGURE 7. Philoria loveridgei holotype NHM 1947.2.19.94 Lamington National Park, Qld. A) dorsal, B) ventral views.

FIGURE 8. Philoria loveridgei in life A) QM J86652 Green Mountains Section, Lamington National Park, Qld; B) No voucher, Mount Mumdjin, Springbrook National Park, Qld; C) AMS R186686 Border Ranges NP, NSW; D) AMS R186687 Border Ranges National Park, NSW; E) AMS R186703 Wollumbin National Park, NSW; F) AMS R186701 Wollumbin National Park, NSW; G) QM J86652 ventral view Lamington National Park, Qld; H) QM J86651 ventral view, Springbrook National Park, Qld; I) AMS R186684 ventral view Border Ranges National Park, NSW; and J) AMS R186701 ventral view Wollumbin National Park, NSW.