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Litoria haematogaster, sp. nov. Red-bellied Treefrog Figs 9–10 https://zoobank.org/ urn:lsid:zoobank.org:act: 561826C8-153C-416D-8E79-2ED52E67B6CD Holotype. SAMA R71702 (FN SJR3205), adult female, Darai Plateau, Gulf Province, Papua New Guinea (7.1295°S, 143.6134°E; 435 m a.s.l.), collected on 29 July 2003 by S. Richards. Diagnosis. Litoria haematogaster sp. nov. is distinguished from all other Litoria by the following unique combination of characters: size moderate (SVL of one female 38.4 mm); dorsum in life mottled pale and darker green with scattered brown spots; vomerine teeth present; fingers extensively webbed; tubercles around vent and along outer edge of tarsus and foot prominent, conical; hidden surfaces of legs and posterior surfaces of belly bright red; purple pigmentation on posterolateral surfaces absent; pigmentation on nictitating membrane restricted to narrow band at dorsal margin; and is genetically diagnosable from L. lisae sp. nov. at 65 sites and from L. majikthise at 40 sites in the 787 base pair alignment of mitochondrial ND4 gene and flanking tRNA (Table 1). Description of holotype. Adult female (Figs 9–10) containing numerous moderately small (~ 1.5 mm diameter) eggs with dark pigment on animal hemisphere. Body moderately slender. Limbs moderately long (TL/SVL 0.56). Head moderately wide (HW/SVL 0.32), slightly less than length (HL/SVL 0.34, HW/HL 0.92). Vomeropalatines forming two short ridges between choanae, without detectable teeth. Tongue large, broadly oval, with distinct posterior notch. Snout rounded in dorsal view with truncate tip (Fig. 10A), steep, near vertical in lateral view (Fig. 10B); canthus rostralis broadly rounded, slightly curved; loreal region steep, slightly concave; lips slightly flared; nostrils much closer to tip of snout than eyes, oriented anterolaterally; internarial distance greater than distance from external naris to eye (EN/IN 0.81); eyes large (EYE/SVL 0.12), prominent, protruding in dorsal and lateral views; pupil horizontal, pigmentation on nictitating membrane restricted to narrow band along dorsal edge. Tympanum large (TYM/SVL 0.06), nearly half diameter of eye (TYM/EYE = 0.47), annulus prominent except dorsal edge obscured by thick, strongly curved supratympanic ridge. Skin of dorsum and dorsal surfaces of limbs finely granular with numerous scattered larger tubercles (Figs 9– 10); throat finely ridged anteriorly, becoming more coarsely ridged posteriorly; a broad band of large, unpigmented tubercles around outer margins of throat; chest and abdomen coarsely granular. Ventral surfaces of limbs mostly smooth except proximal 1/2 distance along posteroventral margins of thighs with numerous large tubercles, those closest to vent largest, with size decreasing distally (Fig. 9B); transverse row of tubercles forming short elevated ridge below vent; ventral to this ridge are four prominent conical tubercles, each measuring ~ 1.2 mm in height; outer edge of tarsus and foot with row of prominent near-conical white tubercles extending from heel to distal subarticular tubercle on Toe 5; heels with seven (left leg) and eleven (right leg) small, pale tubercles; single large but low pale tubercle ventrally on each leg at junction of tibia and tarsus; outer edge of forearm with row of small pale tubercles that grades into more prominent tubercles and short crenulated fold on outer edge of hand where tubercles reach distal subarticular tubercle on Finger 4. Fingers moderately short with expanded terminal discs (3FD/SVL 0.06; 3FD/3FP 2.55) with distinct marginal grooves. Distal subarticular tubercles on fingers 2–4 and proximal subarticular tubercle on Finger 3 strongly bilobed, remaining tubercles unilobed (Fig. 10C); relative lengths of fingers: 3>4>2>1. Finger webbing thick, fleshy, with deep creases forming fleshy ‘nodules’ (Fig. 10C); webbing on inside of Finger 4 reaching to midpoint between distal subarticular tubercle and disc, then extending to disc as fleshy fringe; on outside of Finger 3 reaching to midpoint between distal subarticular tubercle and disc (Fig. 10C), on inside of Finger 3 reaching to base of proximal subarticular tubercle, on outside of Finger 2 reaching to base of disc, and between fingers 1 and 2 reduced to basal fringe. Hand with low, elongate (1.5 mm long) inner, and broad (1.7 mm wide) outer, metacarpal tubercles, the latter partially subdivided into four by deep ridges (Fig. 10C). Toes with expanded terminal discs with terminal grooves, those on toes slightly smaller than on fingers (3FD/4TD 1.15). Distal subarticular tubercles on toes 4–5 bilobed, remainder unilobed; relative lengths of toes 4>5>3>2>1. Toe webbing thick, fleshy, on inside of Toe 5 reaching to base of disc, on both sides of Toe 4 to distal subarticular tubercle, on outside of Toe 3 to base of disc, on inside of Toe 3 halfway to base of distal tubercle, on outside of Toe 2 to base of disc, on inside of Toe 2 halfway to proximal tubercle and on outside of Toe 1 to distal tubercle (Fig. 10D). Foot with prominent, elongate inner metatarsal tubercle, outer tubercle not detectable. Colour in life (Fig. 9). Dorsal and lateral surfaces lime green with darker green flecks and reticulations; scattered pale brown spots across dorsum and on dorsal surfaces of limbs, hands and feet. Ventrally white anteriorly with fine brown stippling on throat, posteriorly abdomen bright red; hidden and ventral surfaces of legs, and axillae, uniform red, except for enlarged ivory tubercles on underside of limbs and around vent; heels white; feet predominantly brown dorsally, hands predominantly green dorsally; outer edges of tarsi and feet, and of forearm, with fine dark brown stippling. Iris pale grey with dark brown veins and pale-yellow rim around pupil. Colour in preservative. In preservative dorsal and lateral surfaces paler and darker blue; dorsal surfaces of fingers 1 and 2 ivory; brown patches on dorsal surfaces of arms still visible, brown pigmentation in life now darker brown; red areas in life pinkish. Ventrally mid-abdomen ivory, ventral surfaces of arms creamy. Throat creamy white with dense brown stippling except on enlarged creamy-white tubercles. Hands ventrally translucent grey blue. Measurements of holotype (in mm).— SVL 38.4; TL 21.5; HL 13.2; HW 12.2; EYE 4.7; TYM 2.2; EN 3.5; IN 4.3; 4TD 2.0; 4TP 1.5; 3FD 2.3; 3FP 0.9; TL/SVL 0.56; HW/HL 0.92; TYM/EYE 0.47; EN/IN 0.81; 3FD/4TD 1.15. Variation. The holotype is the only known specimen. Comparisons. In its moderate size (a female 38.4 mm), slender body, green and brown dorsal colour, and extensively webbed fingers, Litoria haematogaster sp. nov. most closely resembles the following 10 species: L. aplini, L. daraiensis sp. nov., L. gracilis sp. nov., L. iris, L. majikthise, L. nigropunctata, L. richardsi, L. singadanae, L. umarensis and L. verae. Litoria haematogaster sp. nov. is also morphologically similar to the two new species described below and is compared with them in the relevant species accounts. Litoria haematogaster sp. nov. differs from L. aplini by having discrete prominent conical tubercles along the tarsus and foot (vs. a crenulated dermal fold), and hidden surfaces of limbs red (vs. predominantly blue with dark-brown mottling); from L. iris by having hidden surfaces of limbs and posterior surfaces of abdomen red (vs. posterior of thighs blue, red, or yellow, frequently blotched with white or purple, posterior surfaces of abdomen not red), and violet spots in axilla and groin absent (vs. present); from L. nigropunctata by having prominent conical tubercles along the tarsus and foot (vs. a low dermal ridge), and hidden surfaces of limbs and posterior surfaces of abdomen red (vs. yellow); from L. richardsi and L. singadanae in having a smaller (TYM/EYE 0.46 vs. 0.65–0.81), pigmented (vs. substantially transparent) tympanum, and further from L. richardsi in lacking (vs. having) irregular black lines on dorsum and extensive black markings ventrolaterally; from L. umarensis and L. verae in having prominent conical tubercles on tarsus and foot (vs. absent), and further from L. umarensis in having posterior surfaces of thighs red (vs. brown), and dorsum mottled lighter and darker green (vs. normally uniform green); and further from L. verae in lacking (vs. having) small brown spots aligned transversely on dorsum, and having red (vs. orange) on posteroventral surfaces in life. The new species is morphologically most similar to L. majikthise from which it is genetically divergent (dA between the taxa of 0.08) and from which it can most readily be distinguished by having more prominent conical tubercles on the limbs and around the vent (vs. tubercles lower, rounded; e.g., Fig. 9B vs. Fig. 2B in Johnston & Richards (1994)), violet patches in axillae absent (vs. present), and head longer (HL/HW 1.08 vs. 0.81–1.03). Distribution and ecology. Litoria haematogaster sp. nov. is known only from a single location on the Darai Plateau in Gulf Province, southern Papua New Guinea (Fig. 4). The Darai Plateau is an extensive area of limestone karst in the Kikori River Basin and forms part of the Great Papuan Plateau. The holotype was found in primary foothill rainforest where it was perched at a height of 3.5 m on a sapling in a swampy area with small forest pools. It contains moderate-size (1.5 mm diameter) eggs with a distinctly darker animal pole. The reproductive biology of this species is unknown. IUCN Red List status. Litoria haematogaster sp. nov. is known from one locality on the Darai Plateau, which supports an extensive tract of lowland forest in the Kikori River basin. However, logging operations are occurring widely within this catchment so until the species’ distribution, habitat requirements and any potential threats are better documented we recommend that it be listed as Data Deficient by the IUCN. Etymology. From the Greek haema ‘blood’, and ‘gaster’, belly, referring to the bright red colour of the posteroventral surfaces of this species. Molecular divergences. Based on analyses of a 787 base pair alignment from the mitochondrial ND4 gene and flanking tRNA, L. haematogaster sp. nov. is most closely related to L. majikthise (dA between the taxa of 0.08, Table 2). dA between sister species pairs in other groups of Litoria ranges from 0.04 to 0.25 (Donnellan et al. 2021, Rowley et al. 2021). Our molecular data also suggests potential structure across the two sampled populations of L. majikthise (Fig. 18; dA of 0.07), but in the absence of additional acoustic and morphological evidence for differentiation we conservatively consider all these populations to be conspecific at this stage., Published as part of Richards, Stephen J., Donnellan, Stephen C. & Oliver, Paul M., 2023, Five new species of the pelodryadid genus Litoria Tschudi from the southern versant of Papua New Guinea's Central Cordillera, with observations on the diversification of reproductive strategies in Melanesian treefrogs, pp. 151-190 in Zootaxa 5263 (2) on pages 167-170, DOI: 10.11646/zootaxa.5263.2.1, http://zenodo.org/record/7800918, {"references":["Johnston, G. R. & Richards, S. J. (1994) A new species of Litoria (Anura: Hylidae) from New Guinea and a redefinition of Litoria leucova (Tyler, 1968). Memoirs of the Queensland Museum, 37, 273 - 279.","Donnellan, S. C., Catalano, S., Pederson, S., Mitchell, K., Suhendran, A., Price, L. C., Doughty, P. & Richards, S. J. (2021) Revision of the Litoria watjulumensis (Anura: Pelodryadidae) group from the Australian monsoonal tropics, including the resurrection of L. spaldingi. Zootaxa, 4933 (2), 211 - 240. https: // doi. org / 10.11646 / zootaxa. 4933.2.3","Rowley, J., Mahony, M. J., Hines, H. B., Myers, S., Aplin, K. P., Price, L. & Donnellan, S. C. (2021) Two new species from the Litoria rubella species group from eastern Australia. Zootaxa, 5071 (1), 1 - 41. https: // doi. org / 10.11646 / zootaxa. 5071.1.1"]}

Litoria daraiensis, sp. nov. Darai Plateau Treefrog Figs 1–3 https://zoobank.org/ urn:lsid:zoobank.org:act: 0760AD74-4A52-4980-AA2E-46F4C325BA7B Holotype. SAMA R71675 (FN JCUNQ [SJR]3136), adult female, Darai Plateau, Gulf Province, Papua New Guinea (7.1295°S, 143.6134°E; 435 m a.s.l.) on 23 July 2003 by S. Richards. Paratype (n = 1). SAMA R71676 (FN JCUNQ [SJR]3181), adult male, same location and collector details as for holotype except collected on 25 July 2003. Diagnosis. Litoria daraiensis sp. nov. is distinguished from all other Litoria by the following unique combination of characters: size small (SVL of one male 23.9 mm, of one female 30.2 mm); dorsum in life brown with green spots; vomerine teeth absent; finger webbing extensive, reaching nearly to distal subarticular tubercle on inside edge of Finger 4; prominent dermal fold along outer edge of foot; advertisement call a harsh chattering sound lasting 0.35– 0.37 s containing 10–11 distinctly pulsed notes produced at 27–29 notes/s with dominant frequency at 3400–3600 Hz; and is genetically diagnosable from L. gracilis sp. nov. at 46 sites in the 787 base pair alignment of mitochondrial ND4 gene and flanking tRNA (Table 1). Description of holotype. An adult female containing large (1.5–1.8 mm diameter), predominantly yellow eggs. Body slender; limbs moderately long (TL/SVL 0.52); head moderately narrow (HW/SVL 0.30), slightly narrower than length (HL/SVL 0.31, HW/HL 0.96). Vomerine teeth absent. Tongue large, narrowly oval, with distinct posterior notch. Snout broadly rounded in dorsal view (Fig. 1A), steep, bluntly rounded in lateral view (Fig. 1B); canthus rostralis short, strongly curved and broadly rounded; loreal region sloping, slightly concave; lips slightly flared; nostrils much closer to tip of snout than eyes, oriented laterally; internarial distance greater than distance from external naris to eye (EN/IN 0.93); eyes small (EYE/SVL 0.11), but protruding in dorsal and ventral views; pupil horizontal, pigmentation on nictitating membrane restricted to narrow band along dorsal edge; tympanum moderately large (TYM/SVL 0.05), nearly one-half diameter of eye (TYM/EYE = 0.48), ventral 2/3 extremely prominent, dorsal one third obscured by thick, slightly curved supratympanic ridge. Skin of dorsum and dorsal surfaces of limbs finely shagreened; throat ridged anteriorly, becoming coarsely granular posteriorly; chest and abdomen coarsely granular; ventral surfaces of limbs mostly smooth except proximal 2/3 distance along posteroventral margins of thighs with numerous tubercles; dense cluster of prominent white tubercles below vent forms short transverse ridge then tubercles extend distally about 9.5 mm along posteroventral margins of each thigh, losing white colour and becoming translucent distally. A prominent crenulated white dermal fold, nearly unbroken, along outer edge of foot, extending from heel to distal subarticular tubercle on Toe 5; row of small pale tubercles along outer edge of forearm grades into distinct low crenulated fold along outer edge of hand reaching distal subarticular tubercle on Finger 4. Fingers moderately short with expanded terminal discs (3FD/SVL 0.05; 3FD/3FP 1.36) with distinct marginal grooves; distal subarticular tubercles strongly bilobed; proximal tubercle unilobed; relative lengths of fingers 3>4>2>1 (Fig. 1C). Webbing on inside of Finger 4 reaching half-way between distal subarticular tubercle and disc on left hand, but not reaching distal tubercle on right hand, before extending to disc as fleshy fringe; on outside of Finger 3 reaching to base of distal subarticular tubercle (Fig. 1C); on inside of Finger 3 reaching to just above proximal subarticular tubercle and on outside of Finger 2 reaching to base of disc; webbing only a basal fringe between fingers 1 and 2. Hand with low, oval inner and narrow oval outer, metacarpal tubercles. Toes with expanded terminal discs (4TD/4TP 1.18) with terminal grooves, discs on toes smaller than those on fingers (3FD/4TD 1.15), relative lengths of toes 4>5=3>2>1 (Fig. 1D). Webbing on inside of Toe 5 reaches nearly to disc, on both sides of Toe 4 to base of distal subarticular tubercle, on outside of Toe 3 to beyond halfway between distal subarticular tubercle and base of disc, on inside of Toe 3 to base of distal tubercle, on outside of Toe 2 nearly to base of disc, on inside of Toe 2 to proximal tubercle and on outside of Toe 1 to distal tubercle. Foot with prominent narrow, elongate inner metatarsal tubercle, outer tubercle barely detectable (Fig. 1D). Colour in life (Fig. 2A). Dorsum pale, green-tinged ivory with green spots and flecks across dorsal surfaces including limbs; dark-brown pigmentation aggregated into small patches on limbs and lateral surfaces; a large patch of brown pigmentation laterally behind axillae, and another on head where it forms broad ‘V’ shape mark between eyes. Ventral surfaces white anteriorly with clumps of brown pigment near angle of jaws, yellow posteriorly and on hidden surfaces of limbs. Dermal folds on limbs and tubercles on limbs and around vent white. Iris ivory with brown reticulations, narrow yellow rim surrounding pupil. Hands and feet translucent, with dense brown spotting and small patches of green dorsally. Colour in preservative. In preservative dorsal surfaces pale milky blue with scattered darker blue and brown flecks, ventral surfaces of legs pale yellow, tubercles on limbs white, ventral surfaces of hands and feet translucent white. Measurements of holotype (in mm). SVL 30.2; TL 15.8; HL 9.5; HW 9.1; EYE 3.3; TYM 1.6; EN 2.7; IN 2.9; 4TD 1.3; 4TP 1.1; 3FD 1.5; 3FP 1.1; TL/SVL 0.52; HW/HL 0.96; TYM/EYE 0.48; EN/IN 0.93; 3FD/4TD 1.15. Variation. The sole paratype is an adult male. It differs from the holotype in its smaller size (SVL 23.9 vs. 30.2 mm), broader tongue (nearly round without posterior notch vs. narrowly oval with posterior notch), presence of small, pale brown, finely granular nuptial pads on Finger 1, and of long vocal slits located laterally in floor of mouth extending from approximately angle of jaws halfway to front of mouth. Colour of paratype in life (Fig. 2B) differs from holotype in having dorsal and lateral surfaces ivory overlain by dense stippling of dark brown, green spots larger, more extensive on head, tending to coalesce on snout such that snout is substantially green; ‘V’ shaped brown pigment patch on head present but less well defined than on holotype. Brown venation on iris finer than that of holotype, with several small brown pigment patches present (vs. absent) (Fig. 2). Measurements of paratype: SVL 23.9; TL 13.1; HL 8.0; HW 7.1; EYE 3.0; TYM 1.1; EN 2.1; IN 2.4; 4TD 1.0; 4TP 0.8; 3FD 1.2; 3FP 0.8; TL/SVL 0.55; HW/HL 0.89; TYM/EYE 0.37; EN/IN 0.88; 3FD/4TD 1.2. Advertisement call. Two calls produced by the male paratype at an air temperature of 22.6°C were recorded. They comprised a series of 11 and 10 harsh, distinctly pulsed chattering notes lasting 0.35 and 0.37 s (Fig. 3). Note lengths for both calls combined were 0.011 – 0.023 s (mean = 0.018, SD = 0.002, n = 21) and notes contained 4–12 pulses (mean = 9.3, SD = 1.71, n = 20; pulses were insufficiently resolved to count in one note). However only one note contained 4 pulses, the remainder contained 7–11 pulses. Pulse rate within notes was 300–647 pulses/s (mean = 484.65, SD = 71.46, n = 20) and notes within calls were produced at a rate of 27–29 notes/s with dominant frequency at 3400 and 3600 Hz for the two calls. A single call containing 11 notes is illustrated in Fig. 3. Comparisons. In its small size (male SVL 23.9 mm, female 30.2 mm), slender body, green and brown dorsal colour, extensively webbed fingers, and lacking a rostral spike, Litoria daraiensis sp. nov. most closely resembles the following eight species: L. aplini Richards & Donnellan, 2020, L. iris (Tyler, 1962), L. majikthise Johnston & Richards, 1994; L. nigropunctata (Meyer, 1875), L. richardsi Dennis & Cunningham, 2006, L. singadanae Richards, 2005, L. umarensis Günther, 2004 and L. verae Günther, 2004. Richards & Donnellan (2020) provided a comprehensive table comparing morphological and acoustic features among all these species except L. nigropunctata and L. umarensis and we draw upon that table in the comparisons presented below. Litoria daraiensis sp. nov. is also morphologically similar to the other four new species formally described in this paper and is compared with them in the subsequent species accounts. Litoria daraiensis sp. nov. differs from L. aplini by its smaller body size (an adult male 23.9 mm vs. males 31.9–35.1 mm SVL), less prominent dermal fold along outer edges of limbs, hidden surfaces of limbs predominantly yellow (vs. predominantly blue with dark-brown mottling), and advertisement call a series of 10–11 distinctly pulsed chattering notes (vs. advertisement call a short buzz normally followed by 1–7 clicks); from L. iris by its smaller size (male L. iris 25–35 mm SVL), hidden surfaces of limbs predominantly yellow (vs. posterior of thighs blue, red, or yellow, frequently blotched with white or purple), violet spots in axilla and groin absent (vs. present) and advertisement call a series of 10–11 distinctly pulsed chattering notes (vs. advertisement call a series of up to 10 notes of variable length, with long notes preceding or following short notes); from L. majikthise by its smaller size (L. majikthise males 30–35 mm SVL), having posterior surfaces of thighs yellow (vs. uniform red), by lacking a pearl-white post-ocular bar (vs. present), lacking violet patches on posteroventral surfaces of abdomen (vs. present) and advertisement call a series of 10–11 distinctly pulsed chattering notes (vs. advertisement call a single long (0.29– 0.36 s) or short (0.026 –0.080) note); from L. nigropunctata by its smaller size (male SVL 23.9 vs. 27–32 mm; Menzies 1972, 2006; Günther 2004), dermal folds along outer margins of tarsi well developed, distinctly crenulated (vs. dermal folds forming low dermal ridge), and advertisement call a series of 10–11 distinctly pulsed chattering notes (vs. advertisement call comprising an irregular succession of clicks and buzzes; Menzies 1972); from L. richardsi and L. singadanae in having a smaller (TYM/EYE 0.37–0.48 vs. 0.65–0.81), pigmented (vs. substantially transparent) tympanum, and further from L. richardsi in lacking (vs. having) irregular black lines on dorsum and extensive black markings ventrolaterally, and from L. singadanae in lacking (vs. having) extensive area of orange on posteroventral surfaces in life; from L. umarensis in its smaller size (male SVL 23.9 vs. 26.3–30.3 mm SVL), posterior surfaces of thighs yellow (vs. brown), dorsum brown with green spots (vs. normally uniform green) and advertisement call a series of 10–11 distinctly pulsed chattering notes (vs. advertisement call comprising 1–5, normally 2–3, notes); and from L. verae in its smaller size (male SVL 23.9 vs. 33–35 mm SVL), lacking (vs. having) small brown spots aligned transversely on dorsum and lacking (vs. having) extensive area of orange on posteroventral surfaces in life. Distribution and ecology. Litoria daraiensis sp. nov. is currently known from a single location on the Darai Plateau in Gulf Province, southern Papua New Guinea (Fig. 4). The Darai Plateau is an extensive area of limestone karst in the Kikori River Basin and forms part of the Great Papuan Plateau. Both specimens were found in primary foothill rainforest. The female was perched at a height of 1.0 m on foliage over a small forest pool at night. The male was calling at night from a leaf 2.3 m high in a swampy area with a number of small forest pools. The female contains large (1.5–1.8 mm diameter) predominantly yellow eggs that are slightly but distinctly darker on the animal pole. It is likely that this species lays its eggs on leaves overhanging forest pools because the female contains large, pale eggs like those of some other arboreal-breeding species (Menzies 1993), but this has not been confirmed. IUCN Red List status. Litoria daraiensis sp. nov. is known from one locality in the Kikori River Basin (Fig. 4), where it occurs within one of the largest remaining areas of relatively undisturbed lowland forest on the island of New Guinea. However, logging operations are occurring widely within the lower Kikori basin so until the species’ distribution, habitat requirements and any potential threats are better documented we recommend that it be listed as Data Deficient by the IUCN. Etymology. The name daraiensis refers to the type and only known locality of this species, the Darai Plateau in Southern Papua New Guinea. Molecular divergences. Based on analyses of a 787 base pair alignment from the mitochondrial ND4 gene and flanking tRNA, L. daraiensis sp. nov. is most closely related to L. gracilis sp. nov. (average net genetic distance, dA, between the taxa of 0.08, Table 2). dA between sister species pairs in other groups of Litoria ranges from 0.04 to 0.25 (Donnellan et al. 2021, Rowley et al. 2021)., Published as part of Richards, Stephen J., Donnellan, Stephen C. & Oliver, Paul M., 2023, Five new species of the pelodryadid genus Litoria Tschudi from the southern versant of Papua New Guinea's Central Cordillera, with observations on the diversification of reproductive strategies in Melanesian treefrogs, pp. 151-190 in Zootaxa 5263 (2) on pages 154-157, DOI: 10.11646/zootaxa.5263.2.1, http://zenodo.org/record/7800918, {"references":["Richards, S. J. & Donnellan, S. C. (2020) Litoria aplini sp. nov., a new species of treefrog (Pelodryadidae) from Papua New Guinea. Records of the Australian Museum, 72, 323 - 335. https: // doi. org / 10.3853 / j. 2201 - 4349.72.2020.1729","Johnston, G. R. & Richards, S. J. (1994) A new species of Litoria (Anura: Hylidae) from New Guinea and a redefinition of Litoria leucova (Tyler, 1968). Memoirs of the Queensland Museum, 37, 273 - 279.","Dennis, A. J. & Cunningham, M. J. (2006) Litoria richardsi sp. nov., a new treefrog (Anura: Hylidae) from New Guinea. Memoirs of the Queensland Museum, 52, 65 - 69.","Richards, S. J. (2005) A new species of treefrog (Anura: Hylidae: Litoria) from the Huon Peninsula, Papua New Guinea. Zootaxa, 1052 (1), 29 - 39. https: // doi. org / 10.11646 / zootaxa. 1052.1.3","Gunther, R. (2004) Two new treefrog species of the genus Litoria (Anura: Hylidae) from the west of New Guinea. Zoologische Abhandlungen. Staatliches Museum fur Tierkunde in Dresden, 54, 163 - 175.","Menzies, J. I. (1972) Papuan tree frogs of the Litoria nigropunctata group. Herpetologica, 28, 291 - 300.","Menzies, J. (2006) The frogs of New Guinea and the Solomon Islands. Pensoft Publishers, Sofia. [unknown pagination]","Menzies, J. (1993) Systematics of Litoria iris (Anura, Hylidae) and its allies in New-Guinea and a note on sexual dimorphism in the group. Australian Journal of Zoology, 41, 225 - 255. https: // doi. org / 10.1071 / ZO 9930225","Donnellan, S. C., Catalano, S., Pederson, S., Mitchell, K., Suhendran, A., Price, L. C., Doughty, P. & Richards, S. J. (2021) Revision of the Litoria watjulumensis (Anura: Pelodryadidae) group from the Australian monsoonal tropics, including the resurrection of L. spaldingi. Zootaxa, 4933 (2), 211 - 240. https: // doi. org / 10.11646 / zootaxa. 4933.2.3","Rowley, J., Mahony, M. J., Hines, H. B., Myers, S., Aplin, K. P., Price, L. & Donnellan, S. C. (2021) Two new species from the Litoria rubella species group from eastern Australia. Zootaxa, 5071 (1), 1 - 41. https: // doi. org / 10.11646 / zootaxa. 5071.1.1"]}

Litoria lisae, sp. nov. Lisa’s Treefrog Figs 11–13 https://zoobank.org/ urn:lsid:zoobank.org:act: C64BD388-84E0-4288-A6CD-F70DFEABDC35 Holotype. SAMA R71689 (FN SJR15385), Arakubi Quarry, Southern Highlands Province, Papua New Guinea (6.4619°S, 143.2560°E; 1030 m a.s.l.) by S.J. Richards on 16 May 2017. Paratypes. (n = 8, all adult males) SAMA R71688 (SJR14931), same details as holotype but collected on 1 July 2015; R71690 (FN SJR15388), Kilometer Point 107, Iagifu Ridge, Southern Highlands Province, Papua New Guinea (6.4402°S, 143.2240°E; 1,400 m a.s.l.) by C. Dahl and S.J. Richards on 17 May 2015; R60719–60720 (FN SJR2381–2382) and R71686 (FN SJR2127), summit of Iagifu Ridge, Southern Highlands Province, Papua New Guinea (6.4446°S, 143.2193°E; 1360 m a.s.l.) by S.J. Richards on 23 October 2001 (R60719–60720) and 27 May 2002 respectively; R60716 (FN SJR2443), R71687 (FN SJR2445) and PNGNM (FN SJR2444), Gobe Ridge summit, Southern Highlands Province, Papua New Guinea (6.8145°S, 143.7743°E; 830 m a.s.l.) by S. Richards on 29 October 2001. Diagnosis. Litoria lisae sp. nov. is distinguished from all other Litoria by the following unique combination of characters: size small (SVL of males 27.0– 29.7 mm); dorsum in life green with darker green spots; vomerine teeth absent; finger webbing extensive; prominent dermal fold along outer edge of foot; pigmentation on nictitating membrane restricted to narrow band at dorsal margin; advertisement call a single note containing either evenly spaced or discretely clumped pulses; and is genetically diagnosable from L. haematogaster sp. nov. at 65 sites and from L. majikthise at 64 sites in the 787 base pair alignment of mitochondrial ND4 gene and flanking tRNA (Table 1). Description of holotype. Adult male (Fig. 11A) with vocal slits and pale brown nuptial pads. Morphometric data are summarised in Table 4. Body very slender. Limbs long (TL/SVL 0.56). Head narrower than long (HW/SVL 0.28; HL/SV 0.33, HW/HL 0.87). Vomerine teeth absent. Tongue moderately large, oval, with broad posterior notch. Vocal slits short, located laterally in floor of mouth, extending from slightly posterior of angle of jaws to slightly anterior of angle of jaws. Snout narrowly rounded in dorsal view (Fig. 12A), bluntly rounded (nearly truncate) in lateral view (Fig. 12B); canthus rostralis broadly rounded, nearly straight; loreal region steep, slightly concave; lips slightly flared; nostrils much closer to tip of snout than eyes, oriented anterolaterally; internarial distance equal to distance from external naris to eye (EN/IN 1.0); eyes moderately small (EYE/SVL 0.11) but prominent, protruding in dorsal and ventral views; pupil horizontal, pigmentation on nictitating membrane restricted to narrow band along dorsal edge. Tympanum small (TYM/SVL 0.05), slightly less than one half diameter of eye (TYM/EYE = 0.45), distinct except dorsal edge of annulus obscured by thick, strongly curved supratympanic ridge. Skin of dorsum and dorsal surfaces of limbs very finely granular (Fig. 11A). Ventrally throat, chest and abdomen coarsely granular; ventral surfaces of limbs with scattered low tubercles, most prominent on posteroventral margins of thighs. Broken white dermal ridge below vent not extending onto adjacent regions of upper thighs; clump of ~15 prominent white tubercles below dermal ridge; heels with five (right) and seven (left) low, white tubercles; outer edge of foot with series of low, pale tubercles extending from heel nearly to distal subarticular tubercle of Toe 5, those on right tarsus partially fused to form short (3.5 mm) ridge; outer edge of hand with row of small pale tubercles extending from mid-point along forearm to distal subarticular tubercle on Finger 4. Fingers moderately short with expanded terminal discs (3FD/SVL 0.05; 3FD/3FP 1.88) with distinct marginal grooves; subarticular tubercles strongly bilobed; relative lengths of fingers: 3>4>2>1 (Fig. 12C). Webbing on inside of Finger 4 reaching to just above distal subarticular tubercle, on outside of Finger 3 reaching to top of distal subarticular tubercle and continuing to disc as thick flange (Fig. 12C), on inside of Finger 3 reaching to small proximal subarticular tubercle, on outside of Finger 2 reaching to halfway between distal subarticular tubercle and disc, and between fingers 1 and 2 limited to basal fringe. Hand with well-defined narrow inner, and less well defined but detectable outer, metacarpal tubercles. Nuptial pad on Finger 1 pale brown, elongate (1.5 mm long), broader distally than proximally. Toes with expanded terminal discs with terminal grooves, smaller than those on fingers (4TD/SVL 0.04, 3FD/4TD 1.25) (Fig. 12D); relative lengths: 4>5=3>2>1. Webbing on inside of Toe 5 reaches nearly to disc, on both sides of Toe 4 to base of distal subarticular tubercle, on outside of Toe 3 nearly to base of disc, on inside of Toe 3 to halfway between proximal and distal subarticular tubercles, on outside of Toe 2 to slightly more than halfway to distal subarticular tubercle, and between toes 1 and 2 reduced to a basal fringe. Foot with welldefined oval inner metatarsal tubercle, poorly defined outer tubercle (Fig. 12D). Colour in life (Fig. 11A). Dorsal surfaces lime green becoming yellow green laterally and on dorsal surfaces of limbs, green on forelimbs terminating sharply at wrists; dorsal and lateral surfaces with scattered darker green spots; white to yellowish-white bar extends from beneath eye posteriorly to base of forearm; white mid-lateral stripe extends from approximately midway between front and hind limbs to groin; ventrally white anteriorly, except broad pale-brown band around edges of throat, extending onto proximal portions of forelimbs; abdomen yellow posterolaterally and in hidden surfaces of limbs and axillae, yellow in axillae extending approximately halfway along posterior surfaces of forearms; enlarged tubercles on outer edges of limbs and around vent white. Dorsal surfaces of hands and feet stippled with brown spots and blotches, patches of yellowish green dorsally on fingers 3 and 4. Iris ivory with paler ivory rim around pupil and dense dark brown reticulations. Colour in preservative. In preservative dorsal and lateral surfaces mottled darker and lighter blue, brown pigment on throat prominent, much darker than in life; patch of brown pigment midlaterally that was nearly undetectable in life now evident; yellow of posteroventral surfaces and limbs now creamy, pale bar under eye white, more prominent than in life; throat and abdomen creamy with white tubercles. Variation. All of the paratypes are adult males. Morphometric variation is limited (Table 4) but snout shape is variable –the holotype has the most narrowly rounded snout of the type series – in most specimens the tip of the snout is bluntly rounded or in two specimens (SAMA R60716, R71690) near-truncate. All specimens exhibit the distinct white bar beneath the eyes, and in life are yellow on the hidden surfaces of the legs and have extensive areas of yellow on the posterior surfaces of the abdomen (Fig. 11C). However, dorsal and lateral colour in life is highly variable with some specimens being much darker green than the holotype, or with more prominent darker green spots and large brown patches dorsally or laterally (e.g., SAMA R71690, Fig. 11B). In preservative types range from pale to dark blue with scattered darker flecks and blotches dorsally; intensity of dark-brown pigmentation on throat is also variable, ranging from pale brown clumps of pigmentation in SAMA R71686, to intense dark brown band of pigmentation laterally on the throat in the holotype and SAMA R71690. Advertisement call. We recorded 17 calls produced by the holotype at an air temperature of 22.0°C and 13 calls produced by SAMA R71688 at an air temperature of 20.5° C. Calls of both animals were similar so data were combined for analysis but, where relevant, differences are noted. Litoria lisae sp. nov. produces two distinct call types (Fig. 13) at highly variable intervals (inter call interval 0.42– 31.22 s; mean = 6.55 s, SD = 7.22, n = 27), and there appeared to be no pattern to the sequence in which the two call types were produced. Each call type is a single, short note but the rate and pattern of pulse production within notes differ markedly. The first call type is a short ‘buzz’ comprising a rapidly repeated series of evenly spaced pulses, with approximately 0.006 –0.009 s between pulses, while the second call type is a more slowly repeated series of pulses which are ‘clumped’ into discrete groups of 2–3 (Fig. 13) with approximately 0.004 s between pulses within groups and 0.03 s between discrete pulse groups. Although the total length of both call types overlaps extensively the first call type contains more pulses that are produced at a faster rate. We here refer to call type 1 as buzz calls, and call type 2 as chirp calls; they are each described in more detail below. Eleven buzz calls from both animals combined were 0.12– 2.34 s long (mean 0.18, SD 0.03) and contained 10– 22 pulses (mean 16.36, SD 3.69) produced at a rate of 72.3–103.9 pulses/s (mean 87.62, SD 10.53). Although the structure of buzz calls produced by the holotype and SAMA R71688 was extremely similar, the three buzz calls of SAMA R71688 were generally shorter (0.13– 0.17 s vs. 0.16– 0.24 s) and contained fewer pulses (10–13 vs. 16–22) produced at a slower rate (72.29–74.38 vs. 83.76–103.89 pulses/s) than eight buzz calls produced by the holotype. Nineteen chirp calls from both animals combined were 0.11– 0.17 s long (mean 0.15, SD 0.02) and contained 6–12 pulses (mean 8.68, SD 1.82) produced at a rate of 37.26–69.56 pulses/s (mean 52.58, SD 9.71). Again, the structure of chirp calls produced by the holotype and SAMA R71688 was extremely similar and there was extensive overlap in call length between the 10 chirp calls produced by SAMA R71688 (0.107 –0.166 s) and the holotype (0.118 –0.176 s). However, chirp calls of SAMA R71688 contained fewer pulses (6–8 vs. 9–12) produced at a slower rate (37.26–55.04 vs. 52.63–69.56 pulses/s) than the holotype. Whether these call types serve different acoustic functions is not known. Comparisons. In its moderate size (male SVL 27.0– 29.7 mm), slender body, green and brown dorsal colour, extensively webbed fingers, and males lacking a rostral spike, Litoria lisae sp. nov. most closely resembles the following 11 species: L. aplini, L. daraiensis sp. nov., L. gracilis, L. haematogaster sp. nov., L. iris, L. majikthise, L. nigropunctata, L. richardsi, L. singadanae, L. umarensis and L. verae. Litoria lisae sp. nov. is also morphologically similar to the new species described below and is compared with it in that species’ account. Litoria lisae sp. nov. can be distinguished from L. aplini by its smaller body size (adult males 27.0– 29.7 mm vs. males 31.9–35.1 mm SVL), less prominent dermal fold along outer edges of limbs, hidden surfaces of limbs predominantly yellow (vs. predominantly blue with dark-brown mottling), and advertisement call a single buzz or chirp containing either evenly spaced or discretely clumped pulses (vs. advertisement call a short buzz normally followed by 1–7 clicks); from L. daraiensis sp. nov. by its larger size (male SVL> 27.0 mm vs. 23.9 mm), having (vs. lacking) a distinct white bar below the eye, and distinct advertisement call (a single buzz or chirp containing either evenly spaced or discretely clumped pulses vs. 10–11 distinctly pulsed chattering notes); from L. gracilis sp. nov. by having a predominantly green dorsum with darker green and brown spots (vs. predominantly brown with green spots), having (vs. lacking) a distinct white bar beneath the eye, and advertisement call a single buzz or chirp containing either evenly spaced or discretely clumped pulses (vs. call containing 2–5 distinctly pulsed notes); from L. haematogaster sp. nov. by lacking (vs. having) large conical tubercles along outer edge of tarsi and arms, and hidden surfaces of limbs yellow (vs. red); from L. iris by having hidden surfaces of limbs predominantly yellow (vs. posterior of thighs blue, red, or yellow, frequently blotched with white or purple), violet spots in axilla and groin absent (vs. present) and advertisement call a single buzz or chirp containing either evenly spaced or discretely clumped pulses (vs. advertisement call a series of up to 10 notes of variable length, with long notes preceding or following short notes); from L. majikthise by its smaller size (L. majikthise males 30–35 mm SVL), having posterior surfaces of thighs yellow (vs. uniform red), and lacking violet patches on posteroventral surfaces of abdomen (vs. present); from L. nigropunctata by having dermal folds along outer margins of tarsi distinctly crenulated (vs. dermal folds forming low dermal ridge), prominent white bar below eye present (vs. absent) and advertisement call a single buzz or chirp containing either evenly spaced or discretely clumped pulses (vs. advertisement call comprising an irregular succession of clicks and buzzes; Menzies 1972); from L. richardsi and L. singadanae in having a smaller (TYM/ EYE 0.39–0.50 vs. 0.65–0.81), pigmented (vs. substantially transparent) tympanum, and further from L. richardsi in lacking (vs. having) irregular black lines on dorsum and extensive black markings ventrolaterally, and from L. singadanae in lacking (vs. having) extensive area of orange on posteroventral surfaces in life; from L. umarensis in having posterior surfaces of thighs yellow (vs. brown), and dorsum with green and brown spots (vs. normally uniform green); and from L. verae in its smaller size (L. verae males 33–35 mm SVL), lacking (vs. having) small brown spots aligned transversely on dorsum, and having yellow (vs. orange) on posteroventral surfaces in life. Distribution and ecology. Litoria lisae sp. nov. is known from lower montane forest on Gobe Ridge and Iagifu Ridge (including Arakubi) in the Kikori River basin of southern Papua New Guinea (Fig. 4), where they appear to be restricted to limestone karst habitats (Fig. 14). Although several individuals were seen on foliage near a shallow forest pool, calling males were encountered only on low foliage in areas with no evidence of surface water suitable for egg deposition (Fig. 14A) and have been heard vocalising from hidden refuges deep within limestone sinkholes (Fig. 14B). Females have not been detected, and neither eggs nor larvae have been encountered. It is possible that this species breeds in small, subterranean waterbodies within the karst terrain, but this remains to be confirmed. Frog communities at the known localities for L. lisae sp. nov. are otherwise dominated by direct-developing microhylid species. IUCN Red List status. Litoria lisae sp. nov. is known from two areas within the Kikori River basin where extensive forest cover remains. However, logging operations are occurring widely within this catchment so until the species’ distribution, habitat requirements and any potential threats are better documented we recommend that it be listed as Data Deficient by the IUCN. Etymology. The name lisae is an honorific for the senior author’s wife, Lisa Capon, in gratitude for her ongoing support of his research activities. Molecular divergences. Based on analyses of a 787 base pair alignment from the mitochondrial ND4 gene and flanking tRNA L. lisae sp. nov. is most closely related to the clade including L. majikthise and L. haematogaster sp. nov., however it is deeply genetically divergent from both of these taxa (dA between the taxa of 0.13 and 0.14, Table 2). The two sequenced samples of L. lisae sp. nov. from localities 70 kilometres apart also showed some genetic divergence (p-distance 0.024).

Litoria gracilis, sp. nov. Slender Spotted Treefrog Figs 5–7 https://zoobank.org/ urn:lsid:zoobank.org:act: 1E175A08-DD09-44DA-9BF4-D044F0914691 Holotype. SAMA R71697 (SJR2297) Adult male with vocal slits and nuptial pads, small stream adjacent to road ascending Iagifu Ridge nr Moro, Southern Highlands Province, Papua New Guinea (6.3681°S, 143.2232°E; 940 m a.s.l.) by S.J. Richards on 21 October 2001. Paratypes (n = 12). PNGNM (FN SJR8854), SAMA R71673–4 (FN SJR2066–2067), 71677 (FN SJR4930), 71678 (FN SJR8855), 71698 (FN SJR2347) same details as holotype except R71673–4 collected on 15 May 2002, R71677 collected on 30 September 1999, PNGNM (FN8854) and R71678 on 13 November 2004 and R71698 on 22 October 2001; R71699 (FN SJR2361), adult male, Summit of Iagifu Ridge, Southern Highlands Province, Papua New Guinea (6.4446°S, 143.2193°E; 1360 m a.s.l.) by S.J. Richards on 22 October 2001; R71679–71682 (FN SJR10390, 10423–10424, 10438), PNGNM (FN SJR)10422, adult males except R71681 adult female, Juha South, Western Province, Papua New Guinea (5.9018°S, 142.4360°E; 950 m a.s.l.) by S.J. Richards 18–20 February 2008. Diagnosis. Litoria gracilis sp. nov. is distinguished from all other Litoria by the following unique combination of characters: size small (SVL of males 25.1–28.2 mm, of one female 28.6 mm); dorsum in life brown with green spots; vomerine teeth not visible; finger webbing extensive; prominent dermal fold along outer edge of foot; pigmentation on nictitating membrane restricted to narrow band at dorsal margin; advertisement call a short bleating sound lasting 0.13– 0.28 s containing 2–5 distinctly pulsed notes produced at a rate of 7.51–12.00 notes/s with dominant frequency at 3800–4000 Hz; and is genetically diagnosable from L. daraiensis sp. nov. at 46 sites and from L. nigropunctata at 95 sites in the 787 base pair alignment of mitochondrial ND4 gene and flanking tRNA (Table 1). Description of holotype: An adult male with nuptial pads and vocal slits. Morphometric data are summarised in Table 3. Body very slender. Limbs moderately long (TL/SVL 0.52). Head moderately narrow (HW/SVL 0.26), slightly wider than long (HL/SVL 0.25, HL/HW 0.96). Vomerine teeth not visible but detectable by scraping with forceps. Tongue oval with shallow posterior notch. Vocal slits long, located laterally in floor of mouth. Snout broadly rounded with angular tip in dorsal view (Fig. 5A), steeply sloping (near truncate) in lateral view (Fig. 5B); canthus rostralis short, broadly rounded, slightly curved; loreal region steeply sloping, slightly concave; lips slightly flared; nostrils much closer to tip of snout than eyes, oriented laterally; internarial distance greater than distance from external naris to eye (EN/IN 0.82, IN/SVL 0.10, EN/SVL 0.08); eyes moderately small (EYE/SVL 0.11) but prominent, protruding in dorsal and lateral views; pupil horizontal, pigmentation on nictitating membrane restricted to narrow band along dorsal edge. Tympanum small (TYM/SVL 0.04), slightly more than one third diameter of eye (TYM/EYE = 0.39), moderately well-defined except dorsal edge obscured by short supratympanic ridge. Skin of dorsum and dorsal surfaces of limbs finely shagreened, throat finely ridged, abdomen coarsely granular. Ventral surfaces of limbs mostly smooth except numerous low but conspicuous white tubercles extending from vent distally to approximately halfway along postero-ventral margins of thighs; a dense cluster of prominent white tubercles surround vent and clump of small, low, pale tubercles on each heel; low, interrupted, white dermal fold extends along outer edge of tarsus and foot, extending from heel nearly to disc of Toe 5 but reduced to low ridge distally; row of small pale tubercles along distal-most outer edge of forearm grading into low dermal ridge along outer edge of hand, reaching nearly to base of disc on Finger 4. Fingers moderately short with expanded terminal discs (3FD/SVL 0.05) with distinct marginal grooves; distalmost subarticular tubercles on fingers 3 and 4 strongly bilobed, those on fingers 1 and 2 unilobed (Fig. 5C); relative lengths of fingers: 3>4>2>1. Fingers extensively webbed, on Finger 4 webbing extending slightly beyond distal edge of subarticular tubercle at base of penultimate phalanx (distal subarticular tubercle), on outer edge of Finger 3 to about distal subarticular tubercle before continuing as narrow flange to base of disc, on outside of Finger 3 to slightly beyond halfway between distal subarticular tubercle and disc, and between fingers 1 and 2 reduced to basal fringe (Fig. 5C). Inner metacarpal tubercle low, elongate, outer metacarpal tubercle broad, flat. Nuptial pad pale brown, narrow, extending 1.5 mm along outer edge of Finger 1. Toes with relative lengths 4>3=5>2>1. Webbing on inside of Toe 5 and outside of Toe 3 extending nearly to base of discs, on both sides of Toe 4 to distal edge of distalmost subarticular tubercle, on outside of Toe 2 to halfway between distal subarticular tubercle and disc, and on Toe 1 to distal subarticular tubercle. Inner metatarsal tubercle well developed, oval; no outer metatarsal tubercle. Distalmost subarticular tubercles on toes 4 and 5 bilobed, on Toe 3 partially bilobed, and on toes 1 and 2 unilobed (Fig. 5D). Tips of fingers and toes expanded into terminal discs, those of fingers slightly broader than those of toes (3FD/4TD 1.08), all with distinct marginal grooves. Colour in life. Pale creamy brown dorsally and laterally, overlain with numerous small green spots; small patches of brown pigmentation concentrated on dorsal surfaces of tibiae and on outer edges of tarsi; ventrolateral surfaces behind axillae with fine peppering of brown pigment; posterior surface of upper arm largely yellow; dermal folds and large tubercles surrounding vent white. Ventrally white except ventral surfaces of forelegs and upper hindlegs; fine peppering of brown pigment along lateral edges of jaw; bright yellow in groin. Colour in preservative. Dorsal surfaces faded pale blue with darker blue and brown patches, ventral surfaces cream, bright yellow surfaces in life faded to very pale yellow. Variation. Morphometric variation is limited (Table 3). The largest specimen is a female, but sexual dimorphism is slight (SVL of 12 males 25.1–28.2 mm, of one female 28.6 mm). All specimens are very slender (Fig. 6A–C), and in life exhibited a yellow patch on the posterior surfaces of the upper arm, yellow in the hidden surfaces of the legs and on the posterior of the abdomen, and green spots on the dorsum. However, the size and number of green spots was highly variable, as was the density and extent of brown pigmentation and its aggregation into discrete patches of different sizes; some specimens had limited dark-brown pigmentation (e.g., Figs 6A–B) while others exhibited numerous discrete blotches of dark brown on the dorsum (e.g., Fig. 6C) and dorsal surfaces of the limbs. Advertisement call. We analysed nine calls produced by the holotype at an air temperature of 21.0°C and one call produced by SAMA R71680 (Fig. 7) at an air temperature of 21.2° C. The call of L. gracilis sp. nov. is a series of 2–4 (occasionally 5) finely pulsed notes. Calls were 0.13– 0.28 s long (mean = 0.24, SD = 0.06, n = 10) and those of the holotype were produced at 9.0– 44.3 s intervals over a period of 211 s (mean inter-call interval = 25.85 s, SD = 13.66, call rate = 0.04 calls/s). Of the 10 calls analysed in detail two (20%) contained two notes, seven (70%) contained three notes and one (10%) contained four notes (Fig. 7). Eighteen additional calls from an unvouchered male that were not analysed in more detail due to their poorer quality contained 2–5 notes (two notes = 22% of calls, three notes = 39%, four notes = 33%, 5 notes = 6% (a single call)). Notes within calls were produced at a rate of 7.51–12.00 notes/s (mean = 9.52, SD = 1.38, n = 10). Note length for all 10 calls combined was 0.033 –0.057 s (mean = 0.040, SD = 0.005, n = 29) and internote interval within calls was 0.011 –0.105 s (mean = 0.062, SD = 0.025, n = 19). In all calls the last internote interval was longer than preceding intervals (Fig. 7). Notes contained 11–23 pulses (mean = 15.09, SD = 3.06, n = 21) produced at a rate of 263–458 pulses/s (mean = 359.3, SD = 50.9, n = 21). Dominant frequency was 3800–4000 Hz. Comparisons. In its small size (male SVL 25.1–28.2 mm, a female 28.6 mm), slender body, green and brown dorsal colour, extensively webbed fingers, and lacking a rostral spike, Litoria gracilis sp. nov. most closely resembles the following nine species: L. aplini, L. daraiensis sp. nov., L. iris, L. majikthise, L. nigropunctata, L. richardsi, L. singadanae, L. umarensis and L. verae. Litoria gracilis sp. nov. is also morphologically similar to the three new species described below and is compared with them in their respective species accounts. Litoria gracilis sp. nov. differs from L. aplini by its smaller body size (adult males 25.1–28.2 mm vs. males 31.9–35.1 mm SVL), less prominent dermal fold along outer edges of limbs, hidden surfaces of limbs predominantly yellow (vs. predominantly blue with dark brown mottling), and advertisement call comprising 2–5 finely pulsed notes (vs. advertisement call a short buzz normally followed by 1–7 clicks); from L. iris by having hidden surfaces of limbs predominantly yellow (vs. posterior of thighs blue, red, or yellow, frequently blotched with white or purple), violet spots in axilla and groin absent (vs. present) and advertisement call a series of 2–5 finely pulsed notes (vs. advertisement call a series of up to 10 notes of variable length, with long notes preceding or following short notes); from L. majikthise by its smaller size (adult males 25.1–28.2 mm vs. 30–35 mm SVL), having posterior surfaces of thighs yellow (vs. uniform red), by lacking a pearl-white post-ocular bar (vs. present), lacking violet patches on posteroventral surfaces of abdomen (vs. present) and advertisement call a series of 2–5 finely pulsed notes (vs. advertisement call a single long (0.29– 0.36 s) or short (0.026 –0.080) note); from L. nigropunctata by its smaller, though slightly overlapping, size (male SVL 25.1–28.2 vs. 27–32 mm; Menzies 1972, 2006; Günther 2004), dermal folds along outer margins of tarsi distinctly crenulated (vs. dermal folds forming low dermal ridge), advertisement call a series of 2–5 finely pulsed notes (vs. advertisement call comprising an irregular succession of clicks and buzzes; Menzies 1972), and deep genetic divergence ((dA between the taxa of 0.15; see below); from L. richardsi and L. singadanae in having a smaller (TYM/EYE 0.32–0.42 vs. 0.65–0.81), pigmented (vs. substantially transparent) tympanum, and further from L. richardsi in lacking (vs. having) irregular black lines on dorsum and extensive black markings ventrolaterally, and from L. singadanae in lacking (vs. having) extensive area of orange on posteroventral surfaces in life; from L. umarensis in having posterior surfaces of thighs yellow (vs. brown), and dorsum brown with green spots (vs. normally uniform green); and from L. verae in its smaller size (L. verae males 33–35 mm SVL), lacking (vs. having) small brown spots aligned transversely on dorsum, and having yellow (vs. orange) on posteroventral surfaces in life. The new species is most similar to L. daraiensis sp. nov., described above. It can be distinguished from that species by its slightly larger size (male SVL 25.1–28.2 vs. 23.9 mm), different advertisement call (a series of 2–5 finely pulsed notes vs. 10–11 distinctly pulsed chattering notes), and moderately deep genetic divergence (dA between the taxa of 0.08; see below). Distribution and ecology. Litoria gracilis sp. nov. is known from several sites across the southern foothills of Papua New Guinea’s Central Cordillera (Fig. 4) between the upper Strickland River basin in the west and the upper Kikori River basin in the east. The identity of frogs in the upper Fly River basin near the Indonesian border referred to L. nigropunctata by Hyndman & Menzies (1990) requires confirmation, but they may represent this species. We exclude the record of L. cf. nigropunctata from the southern foothills of the Snow Mountains in Papua Province, Indonesia (Richards et al. 2015), pending genetic and acoustic analysis of that population. All records of this species are from foothill rainforest where males call from perches on low foliage adjacent to, or overhanging, small, shallow streams and seepages (Fig. 8). We have observed adult males in close proximity to eggs and embryos attached to leaves over such waterbodies on several occasions (Fig. 6E–F), including at the type locality on Iagifu Ridge. The only other pelodryadid frog observed at this location over many years (1999–2017) is a member of the Litoria genimaculata complex that deposits eggs directly into small streams, and we are confident that these ‘arboreal’ egg clutches belong to L. gracilis sp. nov. This conclusion is supported by the observation that the only known female of this species contains relatively large, yellow eggs typical of species that deposit their eggs on leaves above water (e.g., Menzies 1993). This reproductive strategy is an example of Mode 24 of Haddad & Prado (2005), although the small clear streams into which the tadpoles drop could arguably be considered lotic rather than lentic habitats. IUCN Red List status. Litoria gracilis sp. nov. has a moderately broad distribution across more than 100 km of the southern foothills of Papua New Guinea’s Central Cordillera in an area where extensive tracts of primary forest remain and are likely to do so for the foreseeable future. We therefore recommend that it be assessed as Least Concern by the IUCN Red List. Etymology. Gracilis is a Latin adjective meaning slender, graceful, or gracile, and refers to the slender body form of this species. Molecular divergences. Based on analyses of a 787 base pair alignment from the mitochondrial ND4 gene and flanking tRNA L. gracilis sp. nov. is the sister taxon to L. daraiensis sp. nov. (dA between the taxa of 0.08, Table 2). dA between sister species pairs in other groups of Litoria ranges from 0.04 to 0.25 (Donnellan et al. 2021, Rowley et al. 2021). Samples of L. gracilis sp. nov. from different localities show moderate intraspecific divergence (maximum p-distance 0.052). Litoria gracilis sp. nov. was formerly confounded with L. nigropunctata (e.g., Richards 2003) but it is deeply divergent from topotypic L. nigropunctata from Yapen Island (dA between the taxa of 0.15)., Published as part of Richards, Stephen J., Donnellan, Stephen C. & Oliver, Paul M., 2023, Five new species of the pelodryadid genus Litoria Tschudi from the southern versant of Papua New Guinea's Central Cordillera, with observations on the diversification of reproductive strategies in Melanesian treefrogs, pp. 151-190 in Zootaxa 5263 (2) on pages 161-167, DOI: 10.11646/zootaxa.5263.2.1, http://zenodo.org/record/7800918, {"references":["Menzies, J. I. (1972) Papuan tree frogs of the Litoria nigropunctata group. Herpetologica, 28, 291 - 300.","Menzies, J. (2006) The frogs of New Guinea and the Solomon Islands. Pensoft Publishers, Sofia. [unknown pagination]","Gunther, R. (2004) Two new treefrog species of the genus Litoria (Anura: Hylidae) from the west of New Guinea. Zoologische Abhandlungen. Staatliches Museum fur Tierkunde in Dresden, 54, 163 - 175.","Hyndman, D. & Menzies, J. (1990) Rain forests of the Ok Tedi headwaters, New Guinea: an ecological analysis. Journal of Biogeography, 17, 241 - 273. https: // doi. org / 10.2307 / 2845122","Richards, S. J., Tjaturadi, B., Mumpuni & Puradyatmika, P. (2015) Field guide to frogs of the Mimika region - Papua, Indonesia. PT Freeport Indonesia, Jakarta. [unknown pagination]","Menzies, J. (1993) Systematics of Litoria iris (Anura, Hylidae) and its allies in New-Guinea and a note on sexual dimorphism in the group. Australian Journal of Zoology, 41, 225 - 255. https: // doi. org / 10.1071 / ZO 9930225","Haddad, C. F. B. & Prado, C. P. A. (2005) Reproductive Modes in Frogs and Their Unexpected Diversity in the Atlantic Forest of Brazil. Bioscience, 55, 207 - 217. https: // doi. org / 10.1641 / 0006 - 3568 (2005) 055 [0207: RMIFAT] 2.0. CO; 2","Donnellan, S. C., Catalano, S., Pederson, S., Mitchell, K., Suhendran, A., Price, L. C., Doughty, P. & Richards, S. J. (2021) Revision of the Litoria watjulumensis (Anura: Pelodryadidae) group from the Australian monsoonal tropics, including the resurrection of L. spaldingi. Zootaxa, 4933 (2), 211 - 240. https: // doi. org / 10.11646 / zootaxa. 4933.2.3","Rowley, J., Mahony, M. J., Hines, H. B., Myers, S., Aplin, K. P., Price, L. & Donnellan, S. C. (2021) Two new species from the Litoria rubella species group from eastern Australia. Zootaxa, 5071 (1), 1 - 41. https: // doi. org / 10.11646 / zootaxa. 5071.1.1"]}

Litoria naispela, sp. nov. Crater Mountain Treehole Frog Figs 15–17 https://zoobank.org/ urn:lsid:zoobank.org:act: 876BBBC1-3045-4239-A686-CD995DAFDDD4 Holotype. SAMA R70216 (FN JCUNQ [SJR]3483). Adult male, forest adjacent Herowana Village, Eastern Highlands Province, Papua New Guinea (6.6220°S, 145.1962°E, 1400 m. a.s.l.), by local assistant for S. Richards on 5 December 2003. Paratypes. SAMA R70215 (FN JCUNQ [SJR]3482), R70214 (FN JCUNQ [SJR]3475), adult males, same details as holotype except R70214 collected on 30 November 2003. Referred specimens. SAMA R71597 (FN JCUNQ [SJR]4858), recently metamorphosed juvenile, collected as tadpole on 20 November 1998, preserved on 12 December 1998, locality details as for holotype, collected by S.J. Richards; SAMA R71594–71496 (FN JCUNQ [SJR]3132–3134), subadults, collected as tadpoles by S. Richards on 24–25 November 2001, raised in captivity by A. Mack and D. Wright, preserved on 25 January 2003 by S.J. Richards. Diagnosis. Litoria naispela sp. nov. is distinguished from all other Litoria by the following unique combination of characters: size small (SVL of three males 25.5–25.8 mm); limbs long (TL/SVL 0.60–0.64); tympanum large (TYM/SVL 1.0–1.1, TYM/EYE 0.79–0.82), with predominantly transparent membrane; dorsum in life green with darker green spots, posterior of venter orange; vomerine teeth absent; finger webbing extensive; prominent dermal fold along outer edge of foot; pigmentation on nictitating membrane restricted to narrow band at dorsal margin; and is genetically diagnosable from L. singadanae at 48 sites in the 787 base pair alignment of mitochondrial ND4 gene and flanking tRNA (Table 1). Description of holotype. An adult male (Fig.15A) with vocal slits and nuptial pads. Measurements are presented in Table 5. Body moderately slender. Limbs long (TL/SVL 0.60). Head slightly longer than wide (HW/HL 0.90). Vomerine teeth absent. Vocal slits short, located laterally in floor of mouth, extending anteriorly approximately 2.0– 2.5 mm from corner of jaws. Snout short, truncate, bluntly rounded in dorsal view (Fig. 16A), nearly vertical in lateral view (Fig. 16B); canthus rostralis poorly defined, broadly rounded; loreal region steeply sloping, slightly concave; eyes large (EYE/SV 0.14), protruding distinctly in dorsal and ventral views; pupil horizontal when contracted; pigmentation on nictitating membrane restricted to narrow band along dorsal edge; nares near tip of snout, directed laterally, not visible in dorsal view; tympanum large, conspicuous, (TYM/EYE 0.81), annulus prominent except dorsally where it merges with short, slightly curved supratympanic fold (Fig. 16B). Tympanic membrane transparent except for central connective tissue at attachment of columella, scattered tiny white flecks, and several clumps of black pigment spots (4>2>1. Finger webbing thick, fleshy, on inside of Finger 4 and outside of Finger 2 reaching to base of discs, on both sides of Finger 3 to mid-way between subarticular tubercle at base of penultimate phalanx and disc with fleshy fringe continuing to disc, on outside of Finger 1 reaching to subarticular tubercle at base of penultimate phalanx (Fig. 16C). Subarticular tubercles prominent, those on fingers 3 and 4 distinctly bi-lobed (Fig. 16C), those on Finger 1 and 2 unilobed. Terminal discs moderately large (3FD/SVL 0.06, 3FD/3FP 1.6) with circum-marginal grooves. Nuptial pads narrow, comprising fine pale brown granules, extending ~ 1.8 mm along proximal inner edge of Finger 1. Toes fully webbed except Toe 4, webbed to proximal edge of subarticular tubercle at base of penultimate phalanx; relative lengths 4>5>3>2>1 on left foot, disc of Toe 5 on right foot damaged, so shorter than Toe 3. Subarticular tubercles prominent, distal tubercle on Toe 5 bilobed, on remainder of toes unilobed. Discs moderately large (4TD/SVL 0.06, 4TD/4TP 1.36) with circum-marginal grooves, discs on toes slightly smaller than those on fingers (3FD/4TD 1.07) (Fig. 16C–D). Colour in life (Fig. 15A). Dorsum pale lime green with dark-green spots and small areas of cyan white. Darkgreen pigment patches on dorsum form large ‘Y’-shaped patch between eyes, round patch on one eyelid, and short irregular bars on hindlimbs; very dark-green pigmentation present along outer edge of hand and forearm and forming discrete spots on otherwise transparent tympanic membrane. Outer edge of tarsi white with dark-green blotches. Iris off-white with large brown patch anteriorly and posteriorly. Hidden surfaces of limbs orange, except small yellow patch inside knee; Ventral surfaces of legs orange, orange colour extending anteriorly onto posterior of abdomen, becoming egg-yolk yellow anteriorly; a dark-brown pigment patch laterally on throat just anterior of angle of jaws, remainder of throat with scattered brown pigment spots. Tubercles below vent white. Colour in preservative. Dorsum blue grey with dark-brown and greenish spots; Fingers 1 and 2 white with dark spots; Fingers 3–4 blue grey dorsally with very dark (nearly black) pigment patches; ventrally hands and feet translucent cream, dermal ridges on all limbs and enlarged tubercles around vent white. Variation. In life the two male paratypes were also pale green and both exhibited the prominent broad V shaped inter-ocular marking, which remains evident in preservative; however, they exhibited fewer dark-green markings, and in preservative they are pale-purple grey rather than blue grey, with scattered dark-brown and greenish spots. A recently metamorphosed specimen (SAMA R71597; Fig. 17D) is predominantly brown and white with some patches of brown on the limbs. A subadult specimen (SAMA R71596) raised from a tadpole collected from the treehole illustrated in Fig. 17A is white with medium lime green mottling across head, and light turquoise green mottling and scattered dark-green spots across dorsum; ‘Y’-shaped green marking between eyes is evident. Iris offwhite with narrow brown pigment veins and large brown anterior and posterior pigment patches (Fig. 17E). Comparisons. In its moderate size (male SVL 25.5–25.8 mm), slender body, green and brown dorsal colour, extensively webbed fingers, and males lacking a rostral spike, Litoria naispela sp. nov. most closely resembles the following 12 species: L. aplini, L. daraiensis sp. nov., L. gracilis sp. nov., L. haematogaster sp. nov., L. iris, L. lisae sp. nov., L. majikthise, L. nigropunctata, L. richardsi, L. singadanae, L. umarensis and L. verae. It can be immediately distinguished from all of these species except L. richardsi and L. singadanae by its very large tympanum (TYM/SVL 0.10–0.11, TYM/EYE 0.78–0.80 vs. TYM/SVL L. richardsi by lacking (vs. having) both irregular black lines on dorsum and extensive black markings ventrolaterally; and from L. singadanae by its smaller size (3 adult males SVL 25.5–25.8 vs. 2 adult males 28.7–28.8), longer legs (TL/SVL 0.60–0.64 vs. 0.57 & 0.57), though this difference is less evident when juvenile L. naispela are considered, larger eyes (EYE/SVL 0.13–0.14 vs. 0.11 & 0.11), near absence of pigmentation on throat except patch at angle of jaws (vs. extensive spotting across throat e.g., Fig. 15C vs. 15E), and dense brown pigmentation ventrolaterally absent (vs. present; see Fig. 15C vs. 15E). Litoria singadanae is the closest known relative of L. naispela sp. nov. It is known only from the mountains of the Huon Peninsula, approximately 190 km to the northeast of the type locality (Fig. 4) and these species are separated geographically by the lowlands of the Markham Valley and the high peaks (many to ~ 3,000 m a.s.l.) of the Finisterre/Saruwaged Mountains. Distribution and ecology. Litoria naispela sp. nov. is known only from the vicinity of Herowana Village in Eastern Highland Province, on the southern slopes of Papua New Guinea’s Central Cordillera (Fig. 4). The habitat where the species was encountered is moderately disturbed lower montane rainforest, although large areas around the village had been converted to gardens and coffee plantations. Observations on this species during three separate field expeditions suggest that L. naispela sp. nov. is an obligate treehole breeder. Treeholes utilised by L. naispela sp. nov. were between 1.0 and about 15.0 m above the ground and had openings of at least 10–15cm in diameter (Fig. 17A). Clutch size was between 10– 15 eggs, which were glued to the vertical surfaces of tree trunks about 10–20 cm above the treehole openings (Fig. 17B–C). Embryos developed into fully formed tadpoles within the egg capsule attached to the tree trunk (Fig. 17C) before dropping into the treehole below, usually during heavy rain (S. Richards pers. obs.). This reproductive strategy is an example of Mode 26 of Haddad and Prado (2005) and is shared with its close relative L. richardsi, which has been observed vocalising adjacent to a treehole with egg clutches glued above the opening and free-swimming tadpoles inside (S. Richards, pers. obs.). The reproductive mode of L. singadanae, this species’ closest known relative, remains unknown, but Richards (2005) noted the absence of lentic waterbodies in the area where it was discovered, so it seems highly likely that it has a reproductive strategy like its two relatives. In their contrasting dark brown and white dorsal patterns, recently metamorphosed froglets raised from tadpoles at Herowana bear a striking resemblance in colour and patterning to bird droppings (Fig. 17D). All the treeholes containing L. naispela sp. nov. tadpoles around Herowana were known by local informants to be sources of drinking water frequented by birds. We hypothesis that this species has evolved a colour pattern resembling bird droppings to reduce their detectability to predators when they emerge from treeholes. This mimicry or masquerade behaviour may be convergent on distantly related treefrogs from the Neotropics (Dendropsohus marmoratus Laurenti; Hylidae) and Indochina (Theloderma asperum Boulenger; Rhacophoridae) that have similar colouration and ecology (Toledo & Haddad 2008). IUCN Red-List Status. Litoria naispela sp. nov. is known only from the vicinity of Herowana village, where forest has been extensively cleared for gardens and coffee plantations. However, Herowana is within a protected area, the Crater Mountains Wildlife Management Area, and extensive tracts of primary forest remain in the region. Until the species’ distribution, habitat requirements and any potential threats are better documented we recommend that it be listed as Data Deficient by the IUCN. Etymology. The word naispela is from the Melanesian pidgin meaning ‘pretty’, ‘beautiful’. Molecular divergences. Based on analyses of a 787 base pair alignment from the mitochondrial ND4 gene and flanking tRNA L. naispela sp. nov. is closely related to and strongly supported as the sister taxon to L. singadanae from the Huon Peninsula (dA between the taxa of 0.07, Table 2). dA between sister species pairs in other groups of Litoria ranges from 0.04 to 0.25 (Donnellan et al. 2021, Rowley et al. 2021). The clade comprising these two taxa is in turn allied to L. richardsi, but this taxon is much more divergent from L. naispela sp. nov. (dA of 0.17)., Published as part of Richards, Stephen J., Donnellan, Stephen C. & Oliver, Paul M., 2023, Five new species of the pelodryadid genus Litoria Tschudi from the southern versant of Papua New Guinea's Central Cordillera, with observations on the diversification of reproductive strategies in Melanesian treefrogs, pp. 151-190 in Zootaxa 5263 (2) on pages 177-182, DOI: 10.11646/zootaxa.5263.2.1, http://zenodo.org/record/7800918, {"references":["Haddad, C. F. B. & Prado, C. P. A. (2005) Reproductive Modes in Frogs and Their Unexpected Diversity in the Atlantic Forest of Brazil. Bioscience, 55, 207 - 217. https: // doi. org / 10.1641 / 0006 - 3568 (2005) 055 [0207: RMIFAT] 2.0. CO; 2","Richards, S. J. (2005) A new species of treefrog (Anura: Hylidae: Litoria) from the Huon Peninsula, Papua New Guinea. Zootaxa, 1052 (1), 29 - 39. https: // doi. org / 10.11646 / zootaxa. 1052.1.3","Toledo, L. F. & Haddad, C. F. B. (2008) Colors and some morphological traits as defensive mechanisms in Anurans. International Journal of Zoology, 2009, Article ID 910892. https: // doi. org / 10.1155 / 2009 / 910892","Donnellan, S. C., Catalano, S., Pederson, S., Mitchell, K., Suhendran, A., Price, L. C., Doughty, P. & Richards, S. J. (2021) Revision of the Litoria watjulumensis (Anura: Pelodryadidae) group from the Australian monsoonal tropics, including the resurrection of L. spaldingi. Zootaxa, 4933 (2), 211 - 240. https: // doi. org / 10.11646 / zootaxa. 4933.2.3","Rowley, J., Mahony, M. J., Hines, H. B., Myers, S., Aplin, K. P., Price, L. & Donnellan, S. C. (2021) Two new species from the Litoria rubella species group from eastern Australia. Zootaxa, 5071 (1), 1 - 41. https: // doi. org / 10.11646 / zootaxa. 5071.1.1"]}

Cyrtodactylus hutchinsoni sp. nov. Hutchinson’s Bent-toed Gecko Figs. 1, 2A 3A urn:lsid:zoobank.org:act: 472DCD6D-226C-4690-89A2-E0B0A0337901 Holotype. SAMA R70284 (F# SJR12821), adult female with original tail and alcohol-preserved tissue (ABTC 114857). Collected from an unnamed limestone range approximately 13 km south-east of Hotmin Mission, West Sepik (Sandaun) Province, Papua New Guinea (4.6439ºS, 141.6791ºE; 950 m a.s.l.) by S.J. Richards and D. Ama on 8 December 2009. Diagnosis. Cyrtodactylus hutchinsoni sp. nov. can be distinguished from all other Melanesian and Wallacean Cyrtodactylus by its unique combination of adult size large (SVL up to at least 144.0 mm); body slender; head moderately wide (HW/SVL 0.19); dorsal tubercles flat or at most slightly keeled, rounded, and in approximately 18–20 longitudinal rows at mid-torso; tubercles on ventrolateral folds rounded; tubercles not extending ventrally past ventrolateral folds; tubercles absent on lower jaw and throat; caudal tubercles flat or slightly keeled and at base of tail only; subcaudal scales on original tail not transversely widened; enlarged precloacal and femoral scales forming a continuous series of up to 80 scales that extends to knee; and dorsal colour pattern in life including four pairs of dark-brown dorsal blotches with prominent off-white margins on grey-brown background, and further extensive dark-brown reticulations and small blotchings across head and limbs. Description of the holotype. Adult female of large size (SVL 144.0 mm, TrK 69.6 mm). Head long (HL/SVL 0.25), moderately wide (HW/SVL 0.19), and clearly distinct from neck. Snout rounded in dorsal and lateral profile. Eye to naris distance greater than orbital diameter (EN/OrB 1.42), loreal region slightly inflated, interorbital region and posterodorsal region of snout concave, canthus rostralis rounded. Eyes large (OrB/HL 0.27), pupil vertical with four lobes on each side, supraciliaries extending from anteroventral to posterodorsal edge of orbit, slightly longer at the anterodorsal margin. Ear opening oval, longitudinally widened, bordered above by dorsal skin fold. Rostral roughly pentagonal, about two-thirds as high as wide, with medial suture extending vertically approximately two-thirds ventrally from dorsal edge, bordered dorsally by two flattened supranasals and one much narrower internasal. Nares bordered by first supralabial (point contact), rostral, one enlarged supranasal and enlarged postnasals—five on left and three on right. Supralabials, mostly squarish and ranging from slightly higher than wide to slightly wider than high, 14 to rictus and 10 to midpoint of eye on both sides. Head, temporal and nuchal scales small and granular. Nuchal and temporal regions interspersed with numerous rounded or slightly conical tubercles approximately 2–3 times width of surrounding scales. Enlarged infralabials wider than high, 12 on both sides, bordered by several rows of slightly enlarged scales that grade into small granular gular scales, gular tubercles absent. Mental wider than high, broadly triangular, slightly constricted at point of contact with postmentals. Body moderately gracile and narrower than head; trunk long (Trk/SVL 0.48) with ventrolateral folds. Dorsum with 18–20 middorsal rows (excluding tubercles on ventrolateral folds) of flat or slightly keeled tubercles approximately 3–4 times width of surrounding small granular scales. Tubercles on ventrolateral folds low and slightly conical in dorsal profile, rounded in lateral profile, in series of 35–37. Ventral scales much larger than dorsal scales, increasing in size medially and arranged in approximately 40 rows at midpoint of torso. Precloacal region with patch of enlarged scales beginning approximately five rows from cloaca, continuous with a single row of enlarged femoral scales extending along full length of femur, up to 80 scales across the longest continuous femoral and precloacal series. Limbs slender, forelimbs (FA/SVL 0.15) shorter and less robust than hindlimbs (HDL/SVL 0.18). Lateral and dorsal surfaces of lower forelimbs, and upper and lower hindlimbs with numerous low, rounded tubercles approximately 2–4 times size of surrounding granular scales. Digits long, well developed, inflected at basal interphalangeal joints; subdigital lamellae smooth, rounded, undivided and expanded proximal to digital inflection (6–12–13–14–10 on left manus; 9–11–14–15–13 on left pes); narrow distal to digital inflection (7–9–10–9–11 on left manus; 8–10– 11–10–11 on left pes); large recurved claws sheathed by one dorsal and one ventral scale. Tail original, complete, fixed in recurved position (Fig. 1), 169.0 mm long; scales irregular in shape, medial subcaudal scales twice size of dorsal scales; up to eight indistinct rows of slightly keeled or flat and round dorsal tubercles extending to approximately 10 mm posterior of tail base and approximately 2–3 times size of surrounding scales; a single flat, round lateral tubercle present on each side approximately 5 mm posterior of tail base. Two low, rounded, postcloacal tubercles on each side of tail. Colouration in preservative. Torso dorsally with medium-brown background, overlain by four pairs of irregular dark-brown blotches with light greyish-brown margins, posterior-most pair of blotches much smaller than others (Fig. 1A). Nuchal band dark brown, roughly triangular, extending along posterolateral surfaces of head to posterior edge of eye, thinly divided by a discontinuous light grey-brown medial stripe. Pair of distinct light greybrown stripes extend along posterior margin of nuchal band from shoulder to rictus. A further short grey-brown postorbital stripe present at posterodorsal edge of each eye. Crown of head medium brown with four clusters of irregular dark-brown blotches with indistinct grey-brown margins. Snout light to medium brown with indistinct dark-brown blotching. Supralabials and infralabials buff, with medium-brown speckling. Limbs and digits medium brown dorsally with scattered darker-brown flecks; on hindlimbs and all digits these flecks coalesce to form indistinct bands or blotches. Ventral surfaces of head, torso and limbs buff with scattered light- to medium-brown maculations and flecks, most dense on limbs (Fig. 1B). Dorsal surface of tail medium brown with large irregularly shaped dark-brown blotches, anterior three blotches clearly margined with light grey, posterior blotches with very indistinct margins. Ventral surface of tail medium buff with scattered darker-brown flecks. Postcloacal tubercles light beige with light-brown flecks. Colouration in life. The following description is based on photographs of the holotype taken at night. Pattern in life similar to that in preservative, but more strongly contrasting. Dorsal surfaces of head, neck, trunk and tail greyish brown, lateral surfaces of neck, trunk and posterior of tail grey; these dorsal and lateral base colours are strongly contrasting against darker-brown dorsal and lateral blotches, flecks and mottling (Fig. 2A). Dorsal blotches and nuchal band all have distinct off-white margins and are further mottled with varying tones of brown, including indistinct dark-brown reticulations. Labial scales, postorbital stripes and ventral margins of nuchal band off-white with minimal brown patterning. Dorsal surfaces of limbs grey with distinct to indistinct dark-grey bands or blotches with extensive dark-brown mottling, margins of bands and blotches off-white and indistinct, further dark-brown flecking on limbs. Dorsal surface of tail grey with five large irregular dark-brown blotches or bands with distinct off-white margins followed by two indistinct dark-brown regions without pale margins. Ventral surfaces dark buff with extensive flecking and maculations in varying tones of brown. Ventral surface of tail dark brown with minimal darker flecking, buff anteriorly. Postcloacal tubercles light beige with brown flecks. Iris silvery-grey with extensive chestnut reticulations. Tongue pink. Summary measurements of holotype (mm). SVL 144.0, TL 169.0, OT 169.0, TrK 69.6, HW 27.2, HH 17.1, HL 36.1, EN 13.6, IN 5.0, OrB 9.6, EAR 1.8, FA 21.7, HDL 26.3. Meristics: SUPR 14L/14R to rictus (10L/10R to midpoint of eyes), INFR 12L/12R, DTR 18–20, VENT 40, Toe I LAM (manus) 6 expanded, 7 narrow, Toe 1 LAM (pes) 9 expanded, 8 narrow, Toe IV LAM (manus) 14 expanded, 9 narrow, Toe IV LAM (pes) 15 expanded, 10 narrow, PCTUB 2L/2R. Comparisons. Grismer et al. (2021a) provide definitions and summaries of species groups within Cyrtodactylus and we use these groups in the comparisons below. Cyrtodactylus hutchinsoni sp. nov. can be distinguished from members of the C. arcanus group ( C. arcanus Oliver, Richards & Sistrom and C. manos Oliver, Karkkainen, Rösler & Richards), C. capreoloides group (C. boreoclivus Oliver, Krey, Mumpuni & Richards, C. capreloides Rösler, Richards & Günther, C. medioclivus Oliver, Richards & Sistrom, C. minor Oliver & Richards and C. tanim Nielsen & Oliver) and C. sermowaiensis group (C. atremus Kraus & Weijola, C. crustulus Oliver, Hartman, Turner, Wilde, Austin & Richards and C. sermowaiensis De Rooij) by a combination of large size (SVL> 120 mm versus maximum SVL versus five or more numerous and narrower brown blotches or series of bands, often with extensive brown vermiculations or reticulations). Cyrtodactylus hutchinsoni sp. nov. differs from members of the C. lousiadensis group (C. epiroticus, Kraus, C. klugei Kraus, C. lousiadensis De Vis, C. murua Kraus & Allison, C. robustus Kraus, C. salomonesis Rösler, Richards & Günther, and C. tripartitus Kraus) in lacking transversely widened subcaudal scales (versus present), in having a lower number of dorsal tubercle rows (18–20 versus 20–35), and (with the exception of C. murua) in having a dorsal pattern in life of paired dark-brown blotches on a greyish-brown background (versus alternating and unbroken dark-brown and light-brown transverse bands). Cyrtodactylus hutchinsoni sp. nov. differs from the larger-bodied taxa in the C. novaeguineae group (C. equestris Oliver, Richards, Mumpuni & Rösler, C. irianjayaensis Rösler, C. novaeguineae Schlegel, C. rex Oliver, Richards, Mumpuni & Rösler and C. zugi Oliver, Tjaturadi, Mumpuni, Krey & Richards) in lacking enlarged tubercles ventral to the ventrolateral folds or on the throat (versus present). Cyrtodactylus hutchinsoni sp. nov. differs from the two smaller species in the C. novaeguineae group (C. aaroni Gu ̈nther & Rösler and C. mimikanus Boulenger) in its combination of larger body size (up to at least SVL 144 mm versus maximum SVL of 86.5 mm for C. aaroni and 103 mm for C. mimikanus), in lacking transversely widened subcaudal scales (versus present), and in having a dorsal pattern of a small number of dark-brown irregularly shaped pairs of blotches (versus 7–11 relatively straight and largely unbroken brown bands bordered posteriorly by narrow to wide whitish margins). Cyrtodactylus hutchinsoni sp. nov. is genetically and morphologically part of the C. loriae group. Within this group, populations currently referred to as C. loriae represent a paraphyletic species complex of at least four probable species distributed across eastern New Guinea (Tallowin et al. 2018; Grismer et al. 2021a). In this context, the most critical comparison of the new species is with whichever of these taxa is most likely to be true C. loriae, which was originally described from specimens collected in Moroka district inland from Port Moresby. The following comparisons with C. loriae therefore focus on the type material (n=2), and on two genotyped specimens collected near the type locality (Fane area, Central Province) that are morphologically similar to the types (Fig. 2B). Cyrtodactylus hutchinsoni sp. nov. can be differentiated from these specimens by its larger size (up to at least SVL 144 mm versus maximum SVL of 123 mm), by having a grey-brown (in life) or medium-brown (in preservative) dorsal ground colour (versus yellowish-brown in both life and preservative), by having dark-brown dorsal blotches with strongly contrasting off-white margins (versus very indistinct light-brown blotches without contrasting margins but with the blotches containing two pairs of small dark-brown spots—one anteriorly, one posteriorly—except for the nuchal band, which only has the posterior pair of spots), dorsal surface of head with further dark-brown blotches that strongly contrast against the grey-brown base colour in life (versus no distinct pattern, and overall light yellowish-brown in life), and venter dark buff in life with scattered darker-brown flecks and maculations (versus yellowbrown and unpatterned). From the geographically most proximate known populations of the C. loriae complex from localities in Chimbu Province, including the vicinities of Karimui, Doido, Noru and Yuro Villages (Appendix 1), C. hutchinsoni sp. nov. can be distinguished by its larger size (up to at least SVL 144 mm versus SVL to 128 mm; n = 21) and dark-buff venter in life with dark-brown flecks and maculations (versus buff and unpatterned). Cyrtodactylus hutchinsoni sp. nov. is geographically and genetically most proximate to C. serratus and shares large size with that species. However, it has ventrolateral and caudal tubercles of different shape (low and at most slightly keeled in dorsal profile versus raised and dentate), 18–20 tubercle rows across the dorsum (versus approximately 10–15 rows), and more densely arranged ventrolateral tubercles in the region just superior to the ventrolateral folds (on adults typically more than two rows in the region 5 mm dorsal to the ventrolateral fold versus typically, at most, only one row within the region 5 mm dorsal to the ventrolateral fold), dorsal and lateral tubercle rows on original tails at base only (versus enlarged tubercles extending along the tail in prominent regular whorls) (Fig. 3A–B), and in having light greyish-brown base colouration on the dorsal and lateral surfaces in life (versus usually khaki or darker-brown) (Figs. 2C–D). Genetic differentiation. Based on analyses of mitochondrial ND2 data published elsewhere (Tallowin et al. 2018), C. hutchinsoni sp. nov. (therein listed as Cyrtodactylus cf. serratus) is most closely related to C. serratus (mean p-distances 0.121 –0.129) and is more divergent from C. loriae from near the type locality (p-distance 0.160). Other candidate species identified in the C. loriae group identified by Tallowin et al. (2018) have similar or higher levels of genetic divergence (p-distances 0.127 –0.213). Etymology. Named in honour of Mark Hutchinson, recently retired Curator of Herpetology at the South Australian Museum; first, in recognition of his broad contributions to herpetology spanning ecology, osteology, paleontology, systematics and conservation; and second, and more specifically, acknowledging his support and supervision of both PMO and SJR over the last two decades. Distribution and Habitat. Cyrtodactylus hutchinsoni sp. nov. is known from one location on the northern slopes of Papua New Guinea’s Central Cordillera (Fig. 4), where it was collected from primary upper-hill forest on limestone substrate at an altitude of 950 m a.s.l. (Fig. 5). Given the large extent of apparently suitable habitat in the region, this species probably has a broad distribution along the northern slopes of the Central Cordillera in northwestern Papua New Guinea and adjacent north-eastern Papua Province of Indonesian New Guinea. The holotype is gravid with two well-developed eggs. It was collected from amongst the branches of a large tree that had been felled to make a small clearing for a temporary field camp. This indicates that it is arboreal, like other members of the C. loriae group. Cyrtodactylus sermowaiensis was collected in sympatry with the new species, but was most commonly observed in lower strata of the forest, on shrubs and close to the ground. Cyrtodactylus rex has also been collected from elevations up to at least 425 m a.s.l. in nearby forests (Oliver et al. 2016). Suggested IUCN conservation status. Although this species is only known from the holotype, it comes from an area that is sparsely settled and in which large areas of primary forest remain. The type locality is located on a system of karst ranges that are unsuitable for agriculture and difficult to access. On this basis, we suggest the species be listed as Least Concern., Published as part of Oliver, Paul M., Karkkainen, Denise Taimi & Richards, Stephen J., 2021, A new species of Cyrtodactylus (Squamata: Gekkonidae) from the northern versant of New Guinea's Central Cordillera, pp. 260-270 in Zootaxa 5057 (2) on pages 261-267, DOI: 10.11646/zootaxa.5057.2.6, http://zenodo.org/record/5588104, {"references":["Grismer, L. L., Wood Jr., P. L., Poyarkov, N. A., Le, M. D., Kraus, F., Agarwal, I., Oliver, P. M., Nguyen, S. N., Nguyen, T. Q., Karunarathna, S., Welton, L. J., Stuart, B. L., Luu, V. Q., Bauer, A. M., O'Connell, K. A., Quah, E. S. H., Chan, K. O., Ziegler, T., Ngo, H., Nazarov, R. A., Aowphol, A., Chomdej, S., Suwannapoom, C., Siler, C. D., Anuar, S., Tri, N. V. & Grismer, J. L. (2021 a) Phylogenetic partitioning of the third-largest vertebrate genus in the world, Cyrtodactylus Gray, 1827 (Reptilia; Squamata; Gekkonidae) and its relevance to taxonomy and conservation. Vertebrate Zoology, 71, 101 - 154. https: // doi. org / 10.3897 / vz. 71. e 59307","Tallowin, O. J. S., Tamar, K., Meiri, S., Allison, A., Kraus, F., Richards, S. J. & Oliver, P. M. (2018) Early insularity and subsequent mountain uplift were complementary drivers of diversification in a Melanesian lizard radiation (Gekkonidae: Cyrtodactylus). Molecular Phylogenetics and Evolution, 125, 29 - 39. https: // doi. org / 10.1016 / j. ympev. 2018.03.020","Oliver, P. M., Richards, S. J., Mumpuni & Rosler, H. (2016) The Knight and the King: two new species of giant bent-toed gecko (Cyrtodactylus, Gekkonidae, Squamata) from northern New Guinea, with comments on endemism in the North Papuan Mountains. ZooKeys, 562, 105 - 130. https: // doi. org / 10.3897 / zookeys. 562.6052"]}

Litoria mira Oliver, Rittmeyer, Torkkola, Dahl, Donnellan & Richards, sp. nov. (Chocolate tree frog) (Figs 6 – 8) Litoria caerulea Tyler (1968 in part) Litoria caerulea Dahl, Richards & Novotny (2013) Urn:lsid:zoobank.org:act: FE7D1EA3-C01E-433A-9BC7-E67FA5CFC6EF Holotype SAMA R71114 (field no. SJR15133), adult male, Purari River Basin (7.3518 OS, 145.1904 OE), Gulf Province, Papua New Guinea, 30 ma.s.l., collected by S. Richards and E. Nagombi on 13 February 2016. Paratypes All from Papua New Guinea (n = 10). SAMA R 70446 (field no. SJR 14869), subadult female; SAMA R 71655 (SJR 15015), adult female, upper Strickland River basin (6.2763 OS, 142.1022 OE), 110 m. a.s.l., Western Province, collectedby K. Aplin and S. Richards between 4 and 10 August 2013; SAMA R 71657 R 71658, QM J97086 (SJR 8879 – SJR 8880, 8913) adult males, Wamangu Village (3.7870 OS, 143.6520 OE), East Sepik Province, collected by C. Dahl between 26 and 30 October 2004; SAMA R 71117 (SJR 15085), female, Purari River Basin (7.4910 OS, 145.2189 OE), 130 m a.s.l., Gulf Province, collectedby S. Richards and E. Nagombi 4 February 2016; SAMA R 71116 (SJR 15320), female, Purari River Basin (7.7892 OS, 145.2664 OE), 5 m a.s.l., collected by S. Richards and C. Dahl 9 July 2016; SAMA R 71656 (SJR 15336), male, SAMA R 71115 (SJR 15353), male, PNGNM (SJR 15355), female, Purari River Basin (7.8566 OS, 145.3308 OE), 100 ma.s.l., Gulf Province, collectedby S. Richards and C. Dahl between 15 and 18 July 2016. Diagnosis Litoria mira can be distinguished from all other Litoria by the unique combination of moderately large size (male SUL up to 70.8, female SUL up to 79.6 mm); vomerine teeth present; webbing on hand extending no further than base of penultimate phalanx on fourth finger; limbs without prominent white or yellow lateral folds or ornamentation; lip lacking a white stripe; limbs relatively short and robust (TL/SUL 0.41 – 0.48); parotoid gland present, but not prominent and not fused with prominent gland on top of head; head relatively narrow (HW/ HL 0.91 – 1.0), distinctly tapering from body; dorsal colouration uniformly brown, without white or yellow spots; small violet patch of skin at postero-ventral edge of eye; and ventral surfaces of limbs, torso and throat with moderate to extensive regions densely stippled with dark to medium brown. Description of holotype Adult male with the following measurements (mm): SUL 70.8; TL 29.4; HL 19.9; HW 20.7; ES 8.9; EN 6.5; IN 5.2; EYE 7.3; TYM 4.1; 3FW 3.9; 4TW 3.5. Body robust (Fig. 6), head approximately as wide as long (HL/HW 0.96), narrower than body in dorsal view; snout truncate in dorsal view, rounded in lateral view; labial region very slightly flared; loreal region steeply sloping and slightly concave; canthus rostralis indistinct, weakly curved; nares rounded, much closer to tip of snout than to eyes, oriented laterally, barely visible in dorsal view. Choanae moderate sized, ovoid, separated by a distance ~1.5 times their width; vomerine teeth in rows of 5 or 6 clustered along posterior edge of bifid elevated ridge medial to choanae. Eyes rather small (EYE/SUL 0.10), moderately prominent in dorsal and lateral views, pupil horizontal. Tympanum moderately sized (TYM/SUL 0.58), annulus moderately distinct and raised, dorsal edge obscured by prominent thick downwards-curving supratympanic fold that extends from anterodorsal edge of eye to supra-axillary junction. Skin on dorsal and lateral surfaces of body, head, throat, anterior abdomen, arms and extremities of legs smooth; on posterior abdomen and upper leg coarsely rugose. Arms robust, without skinfolds or ornamentation; fingers with relativelengths III> IV> II> I (Fig. 7 a); discs prominent, with circum-marginal grooves, wider than penultimate phalanx on all digits, subarticular tubercles rounded, indistinct, one on digits Iand two on II – IV, supernumerary tubercles absent, prominent palmar tubercle at base of digit I. Nuptial excrescence in single small, very indistinct, squarish pad at base of digit I (Fig. 7 b). All digits with prominent lateral flanges, webbing basal between Iand II, reaching first subarticular tubercles between digits II and III, and reaching second subarticular tubercles between III and IV. Table 2. Summary of raw morphological data for adult Litoria mira Values shown are means and ranges Legs robust, short (TL/SUL 0.42), without skin folds or ornamentation, relative lengths of toes IV> V> III> II> I; terminal discs prominent and expanded, with circum-marginal grooves, wider than penultimate phalanges on all digits; subarticular tubercles rounded, moderately prominent, one on I – II and V, two on III – IV, prominent ovoid inner metatarsal tubercle at base of digit I. All digits webbed, webbing extends just beyond first subarticular tubercle between I – II, to base of disc on II and to first subarticular tubercle on III, to base of disc on outside of III, and to base of penultimate phalanx on both sides of IV and to base of disc on inside of V. Dermal fringes present on all digits. Dorsal colouration in preservative dark brown over all surfaces with no obvious pattern, upper hindlimbs and digits slightly paler brown than torso. Ventral surfaces pale buff, with large area of dark brown on throat and smaller patches of lighter brown on upper torso, lower forelimbs and across hindlimbs and toes. Variation Mensural variation for the six males, four females and one subadult in the type series is presented in Table 2. Females are larger than males. The smallest female (SAMA R70446, SUL 66.7 mm) does not have fully developed eggs. In colouration, all preserved specimens show the same uniformly dark-brown colouration, with tone varying slightly; larger specimens tend to be slightly paler. Venter always pale buff with brown mottling of varying extent and intensity on the throat, anterior portion of torso, fore and hindlimbs, plantar surfaces of webbing and digits. Appearance in life Generally similar in life as in preservative. Photographs of the holotype (SAMA R70446) show the same uniformly brown dorsal colouration in life (Fig. 6) as in preservative, with the outer edge of the tympanum more sparsely pigmented, and lower lateral regions and extremities of limbs slightly more sparsely pigmented and overall lighter brown. Hidden and exposed surfaces of hindlimbs are uniform brown (Fig. 7 c). SAMA R71655 has a greyish dorsal ground colour in life with fine light-brown mottling and maculations. This specimen is dark brown in preservative. All specimens have a small lightviolet patch of skin at the posteroventral edge of the eye in life, which appears to be diagnostic for the species. Iris silvery grey, with dense black vermiculations and flecks. Comparisons Readily distinguished from all Litoria species outside the L. caerulea group by its combination of large size (adult SUL> 60 mm), limbs without prominent skinfolds or lateral ornamentation, absence of a prominent white lip stripe, nictitating membrane without flecks or reticulations, enlarged paratoid gland above tympanum obvious in life, webbing on fingers never extending beyond penultimate subarticular tubercle, snout rounded in lateral view and plain-brown dorsal colouration without any pattern. Litoria mira differs from L. caerulea in its smaller maximum size (maximum male SUL length 70.8 versus 77.3 mm, maximum female SUL 79.6 versus at least 99.7 mm), generally narrower head (HL/HW mean 0.97, range 0.91 – 1.0 versus mean 0.9, range 0.78 – 0.98); presence of a small violet patch of skin at posterio-ventral edge of the eye in life (versus absent), more extensive brown ventral pigmentation on throat, belly and limbs (versus scattered and generally found on throat region only); and consistently darkbrown dorsal colouration in life (Fig. 6, 8 a, b) and preservative (versus typically green (Fig. 8 c), brownish green or sometimes brown in life, and often blue in preservative). In life, photographs of two individuals also suggest that L. mira has dark-brown hidden surfaces on the thighs (Fig. 7 c) (versus typically pink or yellow in L. caerulea); however, data from more specimens are required to further test the strength of this character. The three other species in the Litoria caerulea group do not occur in New Guinea, and all lack a pale-violet patch of skin at the posterior-ventral edge of the eye. L. mira further differs from L. cavernicola in having a plain dark-brown dorsum (versus usually olive green: Fig. 8 d), smooth to moderately granular dorsal skin (versus strongly granular) and larger size (maximum male SUL length 70.8 versus 51.0 mm, maximum female SUL 79.6 versus 57.0 mm); from L. gilleni by its plain dark-brown dorsum (versus olive green or brown with yellow or white spots: Fig. 8 e); and from L. splendida by its smaller maximum size (maximum male SUL length 70.8 versus 106.0 mm, maximum female SUL 79.6 versus 118.0 mm), much-less-prominent parotoid gland above eye and snout (versus always apparent in both areas on adults), and darkbrown dorsal colouration in life (versus green with scattered white or yellow flecks: Fig. 8 f). Distribution The type series originates from three localities spanning provinces north and south of New Guinea’ s Central Cordillera (Fig. 1). These scattered records, and low genetic divergence between samples, suggest that the species may occur more widely in the difficult-to-access lowland swamp forests across the island of New Guinea. Tyler (1968) lists uncatalogued specimens of L. caerulea in the Australian Museum and Australian National University from near Maprik Village in northern Papua New Guinea that we have not been able to locate. These specimens are also most likely to be L. mira. We have been unable to examine one further specimen of ‘ L. caerulea ’ (RMNH 12370) from Erokwero on the Bird’ s Head Peninsula (Vogelkop) in West Papua Province listed by Tyler (1968). It requires further investigation to confirm which member of the L. caerulea group occurs in far western New Guinea. Habitat and ecology Collecting localities are in both disturbed and undisturbed lowland swamp forest or swampy rainforest (Fig. 2 a, b). The species was typically observed perched on branches within 3 m of the ground. The advertisement call is a deep, rasping bark repeated in long series, ‘crawk, crawk, crawk...’ (Fig. 9), that is indistinguishable to the ear from that of L. caerulea. Five call series from SAMA R71658 recorded at an air temperature of 25 OC contain 33 – 48 calls (=notes) (mean = 40.4, s.d. = 6.58) produced ata rate of 2.6 – 2.8 calls s – 1 (mean = 2.66, s.d. = 0.08) lasting a total of 13 – 17.8 s (mean = 14.96, s.d. = 2.51). The first call of each series is the softest and shortest, and amplitude of calls subsequently increases during each series (Fig. 9 a) except for the last call, which may have a distinctly lower amplitude than the calls preceding it. Calls last 0.10 – 0.17 s, and contain 18 – 23 pulses produced at a rate of ~150 pulses s – 1 . Other Litoria collected in sympatry with L. mira at sites in southern New Guinea included the widespread lowland species L. auae, L. chloristona, L. congenita, L. infrafrenata, L. pygmaea and L. thesaurensis and in northern New Guinea included L. chrisdahli, L. infrafrenata, L. mucro, L. pygmaea and L. thesaurensis. Suggested IUCN red list status Litoria mira is known from widely separated sites in difficultto-access and relatively undisturbed areas of lowland New Guinea. There are large areas of suitable lowland habitat in the intervening areas, and human population density in most of this region remains low. We suggest that the species be listed as Least Concern. Etymology The name mira is the feminine form of the Latin adjective mirum, for surprised or strange, stemming from our surprise in discovering an undescribed member of the predominately Australian L. caerulea group occurring widely across lowland swampy rainforest in New Guinea.

Oliver, Paul M., Rittmeyer, Eric N., Torkkola, Janne, Donnellan, Stephen C., Dah, Chris, Richards, Stephen J. (2021): Multiple trans-Torres Strait colonisations by tree frogs in the Litoria caerulea group, with the description of a new species from New Guinea. Australian Journal of Zoology 68: 25-39, DOI: 10.1071/ZO20071

Oliver, Paul, Günther, Rainer, Tjaturadi, Burhan, Richards, Stephen J. (2021): PAUL OLIVER, RAINER GÜNTHER, BURHAN TJATURADI & STEPHEN J. RICHARDS (2021) A new species of large green treefrog (Litoria, Pelodryadidae) from Papua, Indonesia. Zootaxa, 4903 (1): 117-126. Zootaxa 4948 (4): 599-599, DOI: https://doi.org/10.11646/zootaxa.4948.4.8

Oliver, Paul, Günther, Rainer, Tjaturadi, Burhan, Richards, Stephen J. (2021): A new species of large green treefrog (Litoria, Pelodryadidae) from Papua Indonesia. Zootaxa 4903 (1): 117-126, DOI: https://doi.org/10.11646/zootaxa.4903.1.7

Litoria lubisi sp. nov. Litoria cf. infrafrenata (Richards et al. 2015) Lubis’s Treefrog Figs. 1–2, 4 A. Holotype. MZB Amph. 13.358 (FN SJR 7946). Adult male, calling when collected, 10 km south of Timika town, Papua Province, Indonesia (4.662°S, 136.897°E; 20 m a.s.l.), collected by S.J. Richards and B. Tjaturadi on 8 December 2006. Diagnosis: A large treefrog that differs from other New Guinean Litoria in the following unique combination of characters: body large (adult male SVL to at least 69.1 mm), moderately robust; vomero-palatines large, conspicuous, each with ten small teeth; labial stripe indistinct and not reaching rictus of jaw; dorsum uniformly bright green; parotoid glands absent; scapular region lacking deep skin creases; webbing between Fingers 3 and 4 extending halfway along penultimate phalanges; webbing on feet extending to discs on all toes except Toe 4; discs wide (3FD/SVL 0.071; 4TD/SVL 0.061); nuptial pads crescentic and broader anteriorly than posteriorly; ventral and lateral surfaces yellow suffused with pale blue along skin creases; iris pale reddish brown; upper half of nictitating membrane clear with dark dorsal margin; and advertisement call a single rasping note lasting 0.18– 0.30 s uttered in very long (>5 min) series with irregular inter-note intervals. Description of holotype. Habitus moderately robust, limbs moderately long (TL/SVL 0.54); snout broadly rounded in dorsal aspect, blunt, steeply sloping in lateral aspect. Canthus long, poorly defined, nearly straight; loreal region sloping, distinctly concave; nostrils near tip of snout. Eyes moderately small (EYE/SVL = 0.10) (Fig. 1), nictitating membrane without flecks or lines of pigmentation. Tongue oval with shallow posterior notch; vocal slits long, extending from near angle of jaw half way to front of mouth. Ten small teeth on each of two prominent, slightly postero-medially directed vomero-palatine ridges. Tympanum moderately large (EAR/SVL = 0.082), annulus distinct except postero-dorsal edge obscured by thick, curved, postocular fold that extends from posterior edge of eye to above axillary junction. Skin finely but distinctly granular dorsally, slightly coarser on head. Lateral surfaces with large, flat tubercles; throat finely ridged, belly and ventral surfaces of thighs coarsely tubercular. Prominent dermal ridges along posterior edge of forearms and along shanks and ankles extending to toe 5 (Figs. 1, 2 A–2B). Fingers short, partially webbed, terminal discs wide (3FD/SVL 0.071), with circum-marginal grooves (Fig. 2C). Webbing reaches to half-way between penultimate subarticular tubercle and disc between Fingers 4 and 3, half way between penultimate and third subarticular tubercle on inner side of Finger 3, to disc on outer side of Finger 2, and is restricted to thin basal strip between Fingers 1 and 2 (Fig. 2A). Subarticular tubercles prominent, undivided. Relative lengths of fingers 3>4>2>1. Nuptial excrescences a narrow, unbroken patch extending along approximately half length of Finger 1, with distal half approximately equal in height to minimum width of Finger 1 and proximal half much narrower and tapering proximally, forming an approximate crescent (Fig. 2C). Toes fully webbed, terminal discs moderately wide (4TD/SVL 0.061) with circum-marginal grooves. Webbing reaches discs on Toes 1–3 and 5, and to half-way between penultimate tubercle and disc on Toe 4. Inner metatarsal tubercle and subarticular tubercles prominent (Fig. 2B). Relative lengths of toes 4>5>3>2>1. In life, dorsal and upper-lateral surfaces of torso and head uniformly green without any pattern; tympanic membrane with a distinct unpigmented U-shaped region (Fig. 1); mid-lateral surfaces of torso yellow with extensive pale blue reticulations along skin creases; ventral surfaces of torso and inner surfaces of limbs yellow or yellowish brown (Fig. 2D). Dorsal surfaces of digits light brown or yellowish green, ventral surface of digits yellowish, except discs unpigmented; webbing light purplish-blue, nuptial excrescences light brown. Dermal ridges on, and posterior edges of, forearm and hindlimb yellowish white. Labial stripe indistinct, pale yellow, extending to approximately level with posterior edge of eye. Lower half of nictitating membrane green, upper half clear with a dark-brown dorsal margin. Iris with light-grey base colouration, densely stippled with light brown to give an overall reddish-brown appearance, with scattered sparse darker-brown vermiculations; sclera with thin dark-brown inner ring and slightly wider pale-blue outer ring. Skin above eye with very thin pale yellowish-white band. In preservative, most dorsal and upper lateral surfaces mottled light to medium blue, including torso, head, outer forearms, shanks, Finger 4 and Toe 5; tympanic membrane heavily pigmented apart from broad U-shaped unpigmented mark; inner edge of forearm light buff, digits and webbing buff with faint purplish tinge. Lower lateral surfaces of torso buff with indistinct purplish-blue vermiculations. Ventral surfaces light buff and largely unpatterned, with exception of purplish tinge on webbing and digits, especially on feet. Summary meristic data. SVL 69.1; TL 37.4; HW 26.5; HL 23.7; EYE 7.2; TYM 5.7; IN 5.7; EN 4.7; 3FD 4.9; 3FP 3.1; 4TD 4.2; 4TP 3.8. Advertisement call. The call of Litoria lubisi sp. nov. is a single rasping note, produced in long call series that can last at least five minutes without interruption. Fifty-four calls from a longer series produced by the holotype at an air temperature of 26.5°C have a length of 0.18– 0.30 s (mean = 0.22, SD = 0.031) and were uttered over the course of 66.7 s at a rate of 0.79 calls/s. However, calls within the series tend to be ‘clumped’ into groups of 2–4 (Fig. 3A), with intervals between calls ranging from 0.40– 3.17 s (mean = 1.08, SD = 0.82). Energy increases gradually during each call, reaching maximum amplitude approximately halfway or slightly more than halfway through the call, before declining again (Fig. 3B). Although call length is as much as 0.3 s, most energy is concentrated in the first 0.15– 0.20 s of each call followed by an initially steep but then gradually more shallow decline in amplitude with a long, tapering ‘tail’ of low but detectable energy (e.g. Fig. 3B). Dominant frequency ranges from 1.480 –1.838 kHz (mean = 1.651, SD = 80.794 among calls. Calls are finely pulsed, and pulses are uniformly distributed, but pulses are of differing intensity, including ‘primary’ (strong) and ‘secondary’ (weak) pulses (Köhler et al. 2017). Resolution of the secondary pulses in particular is insufficient to permit pulse counts within calls, particularly in the terminal half, so it has not been possible to calculate pulse rates within calls. To the human ear the call of L. lubisi sp. nov. resembles a rapidly repeated short, harsh, rasping note. Comparisons. Litoria lubisi sp. nov. differs from all other Litoria except taxa in the L. infrafrenata group in having the following set of characters; adult size large (> 55 mm SVL), dorsal colouration uniform and green, paratoid glands not evident pupil horizontal, and webbing not extending to discs on the fingers, but extending discs on most toes. Litoria lubisi sp. nov. (Fig. 4A) differs from the four other species in the L. infrafrenata Group as follows: from L. infrafrenata (Fig. 4B) by having sides yellow suffused with pale blue (versus plain light yellow or off white), pale labial stripe indistinct, ending before posterior edge of lower jaw (versus distinct and extending to tympanum), and advertisement call a single rasping note lasting 0.18– 0.30 s (versus call comprising two notes lasting in total ~ 0.11 s, each note lasting 0.04– 0.045 s; De la Riva et al. 2004); from L. multicolor in its larger size (adult male SVL 69 mm versus SVL of 49.1–55.4 mm, n = 14; Günther 2004), dorsal colouration uniformly green (versus green, grey, brown or yellow, and often patterned with black spots), red spots on venter and thighs absent (versus present), webbing extending to beyond penultimate subarticular tubercle on Fingers 3 and 4 (versus ending at subarticular tubercle on Finger 4 and before subarticular tubercle on Finger 3), and advertisement call consisting of a single rasping note lasting 0.18– 0.30 s (versus ‘wooden rattles’ containing 7–10 notes and lasting 0.53– 0.86 s; Günther 2004); from L purpureolata (Fig. 4C) in having head broader (HW/SVL 0.38 versus 0.33–0.37; n = 10), discs wider (3FD/SVL 0.071 versus 0.054 –0.066; n = 10), venter yellow (versus purple), sides yellow suffused with purple (versus predominately purple), webbing extending distal to penultimate tubercle between Fingers 3 and 4 (versus ending at tubercle), and single-note rasping calls lasting 0.18– 0.30 s at a rate of 0.79 calls/s (versus single-note calls lasting 0.11– 0.17 s uttered at a rate of 1.76 calls/s at the same temperature) (Oliver et al 2007); and from L. sanguinolenta (Fig. 4D) by its larger size (male SVL 69 mm versus maximum male SVL 55 mm; n = 10), latero-ventral regions of torso yellow with light purple reticulations (versus white), webbing extending beyond penultimate subarticular tubercle on Fingers 3 and 4 (versus ending at subarticular tubercle on Finger 4, and before subarticular tubercle on Finger 3) and advertisement call consisting of a single rasping note lasting 0.18– 0.30 s (versus call consisting of 6–8 pulsed notes and lasting 0.31– 0.47 s; De la Riva et al. 2004). Litoria lubisi sp. nov. is also similar in size and build to members of the L. graminea Group but differs from all known species in that group by having webbing on hands not extending to penultimate tubercle on inner side of Finger 3 (versus extending beyond penultimate tubercle). Litoria lubisi sp. nov. further differs from Litoria dux Richards & Oliver, 2006 in having nuptial pads elongate and larger than disc on Finger 1 (versus rounded and half size of disc), iris reddish brown (versus white and reddish), and advertisement calls produced at much shorter intervals (every 0.4– 3.2 s versus every 7–30 s) at similar temperatures, with evenly distributed (versus clumped) pulses (Richards & Oliver 2006); from L. graminea (Boulenger, 1905) by having nuptial pads elongate, larger than disc on Finger 1 (versus rounded and half size of disc), iris reddish brown (versus ‘tan/pale gray with red margin, finely stippled with black’; Kraus 2018), and the sole specimen lacks deep skin creases extending to shoulder (versus usually present); from L. huntorum Richards, Oliver, Dahl & Tjaturadi 2006, by iris reddish brown (versus white with reddish outer rim), size larger (male SVL 69 mm versus 53–60 mm; n = 5), lateral edges of venter and webbing yellow suffused with purplish-blue (versus orange), indistinct labial stripe ending before rictus (versus extending to below tympanum), and advertisement call a short, rasping note lasting 0.18– 0.30 s (versus long guttural grunt lasting 0.7– 0.9 s; Richards et al. 2006); from L. nullicedens Kraus, 2018 by nuptial pads elongate (versus round and small), labial stripe weak, white (versus absent), iris predominately reddish brown (versus reticulated black and dark brown), nictitating membrane without flecking (versus present), and sides, abdomen, and webbing of hands and feet yellow and purple (versus bright orange-red; Kraus 2018); from L. pallidofemora Kraus, 2018 by having a narrower snout (EN/IN = 0.82 versus 0.97–1.04; n = 6, Kraus 2018), lateral edges of venter and webbing yellow suffused with purplish-blue (versus yellow or orange), and iris predominately reddish-brown (versus tan or reddish with a wide black outer margin), and may also be a smaller species (adult male SVL 69.1 mm versus 73.5–79.7 mm; n = 6), although larger sample sizes are required to confirm this; from L. pterodactyla Oliver, Donnellan & Richards, 2019 by its venter and webbing yellow suffused with light blue (versus overall blue), calls shorter (0.18–3.0 versus 0.95– 1.01 s) with uniformly distributed (versus clumped) pulses (S. Richards unpublished data from four calls associated with the L. pterodactyla holotype that are of insufficient quality for more detailed analysis); and from L. sauroni Richards & Oliver, 2006 in having iris reddish-brown (versus with red and black vermiculations), pigment flecks across lower eyelid absent (versus present), nuptial pads elongate (versus small and round), and advertisement calls with much shorter intervals (every 0.4– 3.2 s versus every 7–30 s) at similar temperatures, with evenly distributed (versus clumped) pulses (Richards & Oliver 2006). Distribution and natural history. Litoria lubisi sp. nov. is currently known only from a single locality south of Timika in Papua Province, Indonesia (Fig. 5). The single animal was calling from ~ 8 m high between the fronds of a Sago Palm Metroxylon sagu Rottbøll at night in a densely vegetated, flooded swamp. One other animal was heard calling nearby but the deep water and spiny vegetation prevented its capture. It was producing a vocalisation that to the ear was indistinguishable from that produced by the holotype. This species probably breeds in these lowland rainforest swamps; it was not detected during extensive surveys in non-swamp rainforest habitats near the type locality (S. Richards and B. Tjaturadi pers. obs.). Suggested IUCN Status. Litoria lubisi sp. nov. is only known from a single specimen. There are large areas of suitable lowland habitat across southern Papua Province, and human population density in most of this region remains low. However, until its distribution and potential threats (if any) are better documented, we suggest that the species be assessed at Data Deficient. Etymology: This species is named for Pak Rusdian Lubis in gratitude for his support of our field survey in the PT Freeport Indonesia Contract of Work area. Discussion The description of Litoria lubisi sp. nov. brings the number of recognised species in the L infrafrenata Group to five. Litoria lubisi sp. nov. and L. multicolor have only been reported from their type localities in the western portion of New Guinea (Günther 2004; this paper) (Fig. 5). Litoria purpureolata and L. sanguinolenta occur more widely across Indonesian New Guinea and into far western Papua New Guinea. Litoria infrafrenata has a broad distribution across all of lowland New Guinea, far northern Queensland, Australia, and many nearby islands (Fig. 5). The high proportion of species in the L. infrafrenata Group recorded only from the western portion of New Guinea contrasts with the morphologically and ecologically similar L. graminea Group, for which most diversity has been documented to the east in Papua New Guinea (Kraus 2018; Oliver et al. 2019b). Assuming that these two species groups are monophyletic, this pattern suggests different geographic centres of historical diversification. Members of the L. infrafrenata Group also appear to be more often associated with swampy lowland forest habitats where at least some taxa congregate near the forest floor, while members of the L. graminea Group are associated with lowland or hill forest habitats and are not observed in large numbers below the forest canopy (Richards & Oliver 2006; Kraus 2018; S. Richards, pers. obs). Recent fieldwork and taxonomic investigations have revealed that multiple species of large green treefrogs within both the L. graminea and L. infrafrenata Groups have overlapping ranges across large areas of New Guinea, especially south of the Central Cordillera. To the east in Papua New Guinea this includes three species in the L. graminea Group (L. pallidofemora, L. pterodactyla and L. sauroni) that are relatively similar in body size (Richards & Oliver 2006; Kraus 2018; Oliver et al. 2019b). These are all highly arboreal, and their breeding ecology remains poorly understood. With the description of Litoria lubisi sp. nov., three species in the L. infrafrenata Group are now also known from south of the cordillera in the western (Indonesian) portion of New Guinea. These three species show clearer differentiation in size than species in the L. graminea Group, providing at least one potential axis of ecological differentiation between these putatively related species. Furthermore, there is at least some evidence of habitat differentiation. Litoria lubisi sp. nov. has only been observed in flooded lowland swamp habitats dominated by Sago palm and Pandanus species. This habitat type is likely to be under-surveyed because it is often flooded, has few roads and supports large populations of both mosquitos and crocodiles, making it extremely difficult to access. In contrast L. sanguinolenta breeds in shallow pools in flooded lowland rainforest (Richards pers. obs.), while the extremely large L. infrafrenata is widespread and common in both natural and disturbed habitats.

Oedura bella Northern Territory: Type series. AMS R53437 ���8, 37 km north of MacArthur River base camp on Borroloola Road (16.10��S, 136.12��E); AMS R 53467, Caranbirini Water Hole, 21 km north of MacArthur River base camp (16.22��S, 136.15��E); AMS R 53643, Glyde River, 10 km east of MacArthur River base camp (16.43��S, 136.17��E); AMS R 53782, 37 km north of MacArthur River Camp (16.10��S, 136.12��E); NTM R 19310, Borroloola (16.07��S, 136.30��E); SAMA R 34188, McArthur River Station (16.67��S, 135.85��E); NTM R 37983, Mallapunyah Station (16.97��S, 135.80��E); NTM R 37389, Caranbirini Conservation Reserve (16.27��S, 136.08��E); NTM R 37395, Echo Gorge camp rocks (17.17��S, 137.71��E); NTM R 38035, China Wall Gorge, Nicholson River (17.85��S, 137.42��E); NTM R 21288, Musselbrook Reserve, turnoff to Murray Spring (18.61��S, 138.08��E); NTM R 37947, Mallapunyah Station (16.97��S, 135.80��E); NTM R 37383, Southern Lost City, Limmen National Park (15.63��S, 135.46��E). Queensland: QM J94016 (holotype), 10 km south of Mt Isa on Boulia Road, Queensland, Australia (20.8617��S, 139.4617��E); NTM R 21288, Musselbrook Reservoir (18.61��S, 138.08��E); SAMA R34208 ���9, SAMA R 35425, Lawn Hill NP (18.58��S, 138.50��E); QM J52748, Lawn Hill Station, Century Project Site (18.75��S, 138.58��E); QM J74927, Hells Gate (17.47��S, 138.37��E); QM J75207 ���8, Lawn Hill (18.71��S, 138.48)., Published as part of Oliver, Paul M., Jolly, Chris J., Skipwith, Phillip L., Tedeschi, Leonardo G. & Gillespie, Graeme R., 2020, A new velvet gecko (Oedura: Diplodactylidae) from Groote Eylandt, Northern Territory, pp. 438-450 in Zootaxa 4779 (3) on page 449, DOI: 10.11646/zootaxa.4779.3.10, http://zenodo.org/record/3835377

Oedura marmorata Northern Territory: BMNH xxii.2b, (lectotype), Coburg Peninsula (no exact co-ordinates); NTM R 13295, Victoria River Gregory National Park (15.94��S, 130.50��E); NTM R19029 ���31, North Marchinbar Island (11.28��S, 136.63��E); NTM R 21895, Low Hill, Roper River (14.65��S, 134.35��E); NTM R 22444, Limmen Gate National Park (15.78��S, 135.33��E); NTM R 33814, Bauhinia Downs Station (15.93��S, 135.32��E); NTM R 36709, Fish River Station (14.07��S, 130.79��E); NTM R 36746, Wongalara (14.13��S, 134.34��E); NTM R 22774, Astell Island (11.87��S, 136.42��E); NTM R 36717, Wongalara Sanctuary (14.11��S, 134.22��E); NTM R 36731, Jump-up site, Wongalara Sanctuary (14.15��S, 134.16��E); NTM R 36801, Donydji outstation (12.84��S, 135.47��E); NTM R36830 ���1, Stuart Hwy, 11km N of Pine Creek (13.76��S, 131.76��E); NTM R36837 ���9, Umbrawarra Gorge Road, near Pine Creek (13.97��S, 131.70��E); NTM R36858 ���9, ~ 40km east of Stuart Hwy on Kakadu Hwy (13.68��S, 132.05��E); NTM R36862 ���3, Hayes Creek Wayside Inn (13.59��S, 131.46��E); NTM R36889 ���90, ridge on Central Arnhem Road, just before Barunga (14.51��S, 132.85��E); NTM R36891 ���2, 5 km north of Katherine on Stuart Hwy (14.44��S, 132.24��E); NTM R36893 ���5, road into Eva Valley, south of Manyallaluk (14.46��S, 132.78��E); NTM R 36897, Umbrawarra Gorge (13.97��S, 131.70��E); NTM R 36898, Hayes Creek Wayside Inn (13.59��S, 131.46��E); NTM R 36899, ~ 40km east of Stuart Hwy on Kakadu Hwy (14.68��S, 133.05��E); NTM R36909 ���10, near Manton River Dam, just off Stuart Hwy (12.87��S, 131.14��E); NTM R 37058, Robin Falls, south of Adelaide River township (13.37��S, 131.14��E); NTM R 37059, Daly River Road, near Wooliana Road turnoff (13.72��S, 130.71��E); NTM R 37061, ~ 90 km SW of Katherine, on Victoria Hwy, just north of Flora River Reserve turnoff (15.03��S, 131.78��E); NTM R37067, R37069, ~ 30 km SW of Katherine, on Victoria Hwy (14.67��S, 132.09��E); NTM R 37070, ~ 25 km SW of Katherine, on Victoria Hwy (14.65��S, 132.12��E); NTM R 37074, ~ 50 km N Katherine, on Stuart Hwy (14.20��S, 132.04��E); NTM R 37081, Edith Falls Road, ~ 5 km from Stuart Hwy (14.19��S, 132.07��E); NTM R 37091, Dorat Road, ~ 6 km from Stuart Hwy (13.29��S, 131.12��E); NTM R37147 ���8, 10 km N of Katherine (14.41��S, 132.20��E); NTM R 37149, 2 km N of Katherine (14.46��S, 132.25��E); NTM R 37277, Limestone Gorge, N of Bullita (16.05��S, 132.38��E); NTM R 37330, Blackfellow Creek (13.54��S, 130.83��E); NTM R 37355, Victoria River Region Escarpment Walk (15.61��S, 131.12��E); NTM R 37357, Hayes Creek (13.59��S, 131.46��E); NTM R 37368, 2 km south of Tomato Island (14.76��S, 134.68��E); NTM R 37370, Ngukurr, Roper River (14.65��S, 134.78��E); NTM R37415 ���16, Limestone outcrop at Charles Darwin University Katherine (14.24��S, 132.9��E); NTM R37424 ���5, unnamed Gorge in scarp country, 8 km north of Katherine (14.40��S, 132.14��E); NTM R 37430, Limestone country along Stuart Hwy, south east of Katherine (14.50��S, 132.42��E); NTM R 37431, south of Stuart Hwy, east of Katherine (14.50��S, 132.42��E); NTM R37432 ���33, Stuart Highway, just north of Katherine (14.41��S, 132.20��E); NTM R 37435, South of Hayes Creek (13.61��S, 131.53��E); NTM R37689 ���90 (15.02��S, 130.47��E)., Published as part of Oliver, Paul M., Jolly, Chris J., Skipwith, Phillip L., Tedeschi, Leonardo G. & Gillespie, Graeme R., 2020, A new velvet gecko (Oedura: Diplodactylidae) from Groote Eylandt, Northern Territory, pp. 438-450 in Zootaxa 4779 (3) on pages 449-450, DOI: 10.11646/zootaxa.4779.3.10, http://zenodo.org/record/3835377

Oedura gemmata Northern Territory: NMV D72584, NMV D72598 ���600; NMV D72621, Near Kikiyown (also called ���Kikikyon���) (12.19��S, 133.81��E); NTM R 27383, Oenpelli (12.38��S, 133.02��E); NTM R 27599, Kikiyown (12.19��S, 133.81��E); NTM R34986 ���7, Mountain Valley Station (13.98��S, 133.18��E); NTM R 35680, Kakadu National Park (13.86��S, 132.98��E); NTM R 35753, Oenpelli Reservoir (12.38��S, 133.07��E); QM J83987 ���8, Kolorbidahdah (12.65��S, 134.29��E); NTM R 09864, Buffalo Camp, Baroalba Creek (12.8��S, 132.82��E); NTM R 13414, Gimbat Station, Coronation Hill Area (13.58��S, 132.58��E); NTM R 27599, R 27600, R 27592, Kikikyaw (12.19��S, 133.81��E); NTM R 25362, R 25365, R 25367, Mount Brockman (12.75��S, 132.92��E); NTM R34986 ���7, Mountain Valley Station (13.98��S, 133.18��E); NTM R 02506, Mt Brockman (12.75��S, 132.56��E); NTM R 27383, Oenpelli (12.38��S, 133.02��E); NTM R 37315, Oenpelli (12.39��S, 133.03��E); NTM R09862 ���3, Radon Creek, (12.75��S, 132.93��E); NTM R 22274, Red Fin Creek, 8 Km W of Mary River Crossing (12.75��S, 131.47��E); NTM R 35753, Reservoir, Oenpelli (12.38��S, 133.07��E); NTM R 09877, 5km west of South Alligator River (12.68��S, 132.45��E); NTM R 08880, creek above Gunlom Falls (13.97��S, 132.52��E)., Published as part of Oliver, Paul M., Jolly, Chris J., Skipwith, Phillip L., Tedeschi, Leonardo G. & Gillespie, Graeme R., 2020, A new velvet gecko (Oedura: Diplodactylidae) from Groote Eylandt, Northern Territory, pp. 438-450 in Zootaxa 4779 (3) on page 449, DOI: 10.11646/zootaxa.4779.3.10, http://zenodo.org/record/3835377

Oedura nesos sp. nov. Groote Eylandt Marbled Velvet Gecko Figs. 1–4 Oedura bella (in part) Oliver & Doughty, 2016 Oedura “Marmorata North 6” Rosauer et al. 2016 Holotype. NTM R 38578, adult female with original tail, and liver samples stored in ethanol, from Cave Paintings Recreation Area (13.97322°S, 136.50316°E) Groote Eylandt, Northern Territory, Australia, collected by Graeme Gillespie and Jaime Heiniger on 14 November 2016. Paratypes. (n=12) All from Groote Eylandt, Northern Territory, Australia. NTM R 38576 & R 38582 same locality and date as holotype; NTM R 38579 – R 38581 from sandstone outcrop 700 m north of Alyangula (13.8412°S, 136.42305°E) collected 16 November 2016; NTM R 38583 from sandstone outcrop adjacent to Umbakumba Rd (13.93050°S, 136.4945°E) collected 17 November 2016; NTM R 38577 from sandstone outcrop adjacent to Umbakumba Rd (13.8927°S, 136.5112°E) collected 17 November 2016; AMS R138727–8, Groote Eylandt (13.8299°S, 136.4200°E); NTM R7494 & R7495, Umbakumba Rd (13.8799°S, 136.5000°E); NTM R7541, Ayakamindadina (13.97°S, 136.60°E). Referred material. Tissue samples, stored at the Australian National University, Canberra. CCM4191, CCM4195, sandstone escarpment, Central Groote Eylandt (13.9736°S, 136.6033°E). Diagnosis. Differs from all Oedura in the following combination of characters: moderate size (to 85 mm SVL); head moderately wide (HW/SVL 0.18–0.20) and flattened (HD/SVL 0.07–0.12); tail short (TL/SVL original 0.52–0.62, regrown 0.45–0.54), transversely flattened (shallower than body), much narrower than head (TW/HW 0.59–0.77) and tapering gradually to tip; rostral 50% or less divided; postcloacal spurs 2 or 1 on each side (mode 2); 16 precloacal pores in adult males; head colouration dark brown with pale whitish-yellow supraoculars and canthal stripe absent or indistinct; and dorsal colouration of adults including five moderately well-defined whitish bands (including nuchal band), usually with greyish central shading, alternating with wider regions of extensive yellow splotching on a dark brown background. Description. A medium-sized (adult SVL 70.0–85.0 mm) and robust Oedura (Trk/SVL 0.37–0.48); head moderately wide (HW/SVL 0.18–0.20) and flat (HD/SVL 0.07–0.12), although some variation in head width and depth likely attributable to body condition and quality of preservation. Rostral 50–30% divided, bordered dorsally by two nasals, nasals bordered dorsally by two supranasals and 1–2 (mode 2) similar sized intervening scales. Supralabials 9–10 to midpoint of eye, 11–12 in total; infralabials 9–11. Forelimbs and hindlimbs of moderate length (FA/SVL 0.12–0.14, TA/SVL 0.14–0.17). Subdigital lamellae moderately expanded and prominent, 7–8 under third finger, 8–9 under third toe. Distal lamellae divided on all digits, proximal lamellae undivided. Apical lamellae on terminal scansors separated from more proximal pairs. Subapical lamellae series on fingers 2–5 slightly flared and equal to or narrower than width of terminal scansors on fingers. Subapical lamellae series on toes 2–5 flared, equal to or wider than terminal scansors on toes. Postcloacal spurs present and relatively distinct, mode of 2 and 2 (7 out of 11 adults), more rarely 1 and 2 (4 out of 11 adults). Adult males (n=2) with 16 precloacal pores, in a broken series separated medially by 2 non-pore bearing scales. Original tail short (TL/SVL 0.52–0.62), moderately narrow (TW/HW 0.59–0.77) and strongly depressed in cross section, tapering relatively gradually to tip and with no obvious ventral groove, some variation in width and depth likely attributable to body condition and quality of preservation. Caudal scales on original tail arranged in regular transverse rows of irregularly shaped and sized scales. Dorsal and ventral caudal scales of similar size. Regrown tails short (TL/SVL 0.45–0.54), narrow (TW/HW 0.64–0.73) and strongly depressed in cross section, tapering relatively gradually to tip and with no obvious ventral groove. Some variation in width and depth likely attributable body condition and quality of preservation. Caudal scales on regrown tail arranged in irregular transverse rows of variable sized scales, ventral scales tending to be larger than caudal scales. In preservative, base coloration of dorsum on adults is purplish brown, with five distinct whitish to buff transverse dorsal bands extending from nape to groin. Extensive further light buff blotching and flecking usually present between the light bands and elsewhere on the dorsal and lateral surfaces of the head, torso and limbs, with blotches tending to be quite large, usually more than four scales wide. Light and brown pigmentation on head usually broken and never forming a distinct continuous canthal stripe. Venter plain light buff, with variable amounts of tiny brown maculations on the terminal lamellae, around the infralabials, and sometimes extending onto the lateral regions of the throat. Original tails with four light buff dorsal bands, separated by slightly wider purplish-brown regions with moderate to very extensive areas of buff blotching and flecking. Subcaudal surfaces mostly light buff, with varying amounts of light brown, esepecially around edges, and sometimes forming indistinct light transverse sudcaudal bands. Regrown tails dark brown with extensive and variable light buff splotching and spotting but no regular pattern. Hatchlings and small juveniles lack light buff flecking and generally have a very simple pattern of five sharplydefined light bands on the nape and body, and 4–5 light bands on the tail, separated by broad plain dark brown regions. As specimens grow, the sharp definition of light and dark bands decreases, and the amount of light flecking and blotching across the body increases. In life, the basic adult pattern and coloration consists of three clear and contrasting colouration elements (Fig. 2 A–D, F): background dorsal colouration of dark purplish brown; extensive yellow blotching and spotting overlaying dark-brown regions; and whitish transverse bands with small areas of underlying dark purplish brown visible. Summary measurements of the adults in type series (n=11). Quantitative measurements (mean and range) in mm: SVL 80 (70–85); HW 16.4 (14.5–17.7); HD 7.6 (5.4–9.7); HL 20.9 (19.3–22.0); EN 7.1 (6.6–7.5); IN 3.1 (2.8–3.4); IO 7.2 (6.6–7.9); EYE 5.3 (4.6–5.8); TrK 34.7 (28.0–40.6); Crus 10.4 (8.1–11.4); Tibia 11.8 (9.7–13.5); 3FW 1.8 (1.5–1.9); 3TW 2.1 (1.8–2.4); TL (original) 41.4 (36.0–44.8); TW (original) 10.2 (8.6–11.5); TD (original) 5.3 (3.2–6.9); TL (regrown) 41.4 (36.0–44.8); TW (regrown) 11.6 (9.8–12.7); and TD (regrown) 4.0 (3.5–5.3). Scale counts: SuL 9.5 (9–10); InF 10.6 (9–11); CS 1.8 (1–2); FLAM3 7.7 (7–8); and TLAM3 8.1 (8–9). Particulars of the holotype. Adult female with original tail. Quantitative measurements in mm: SVL 85.0; HW 11.7; HD 7.8; HL 22.0; EN 7.5; IN 3.2; IO 7.5; EYE 5.4; TrK 33.7; Crus 10.6; Tibia 11.6; TL 43.0; TW 12.6; TD 3.8; 3FW 1.9; and 3TW 2.2. Scale counts: SuL 10 (12 to rictus of jaw); InF 10; CS 2/2; FLAM3 7; and TLAM3 8. Comparisons. Oedura nesos sp. nov. differs from its three closest relatives in the Oedura marmorata complex from northern Australia in aspects of tail morphology (both original and regrown) and colour pattern (Figs. 2–4). Oedura nesos sp. nov. was formerly confounded with Oedura bella, from which it differs in having a dorsal pattern including alternating transverse bands of white then regions of brown heavily blotched with yellow (versus yellow dots coalescing into distinct to indistinct yellow bands; Figs. 3–4), and in having no or a very indistinct and wavy canthal stripe (versus clearly defined, narrow, straight and continuous or near continuous canthal stripe; Figs. 3–4). Oedura nesos sp. nov. differs from Oedura gemmata in its smaller size (SVL 70–85 mm versus 84–103 mm), in having a tail (original or regrown) much narrower than the head (TW/HW 0.59–0.77 versus 0.75–1.08), and in having prominent white bands and large yellow splotches on the dorsum (versus smaller yellow or white spots only). Oedura nesos sp. nov. differs from Oedura marmorata in its smaller size (SVL 70–85 mm versus 77–97 mm), and in having a tail (original or regrown) that is much narrower than the head in dorsal perspective versus nearly as wide to wider than head (TW/HW 0.59–0.77 versus 0.76–1.2) and gradually tapering to the tip (versus sharply tapering to attenuated tip; Fig. 2–4). It can be further differentiated from geographically proximate populations of Oedura marmorata in eastern Arnhem Land by having yellowish-white supraoculars in life versus bright yellow supraoculars in life (Fig. 2), and tending to have a tail that is highly depressed in cross-section and shallower than body versus a tail that is comparatively rounded in cross-section and as deep or deeper than the body (Fig. 3–4). Hatchling and small juvenile Oedura nesos sp. nov. are easily distinguished from juveniles of its two relatives in the Oedura marmorata complex with a similar colour pattern (Oedura bella and Oedura marmorata) from northern Australia by having white bands (versus yellow; Fig. 3). Of the other species that are more phylogenetically divergent and geographically distant, Oedura nesos sp. nov. differs from Oedura cincta De Vis, 1888, Oedura fimbria Oliver & Doughty, 2016, and Oedura luritja Oliver & McDonald, 2016 (three arid-zone taxa also formerly included in Oedura marmorata) in its smaller size (SVL 70–85 mm versus 78–108 mm O. cincta, 71–104 mm O. fimbria, 85–99 mm O. luritja), and shorter tail (original tail TL/ SVL 0.52–0.62 versus 0.61–0.87) which is also distinctly flattened in cross section (versus rounded). Oedura nesos sp. nov. differs from the three Oedura endemic to the Kimberley region of north-western Australia as follows: from Oedura gracilis King, 1984 by its short and transversely flattened tail (versus very long tail that approaches length of body) and higher number of paired lamellae on the digits (more than 4 versus less than 4) arranged in slightly flared series (versus tapering); and from both Oedura filicipoda King, 1984 and Oedura murrumanu Oliver, Laver, Melville & Doughty, 2014 in its smaller size (max SVL 85 mm versus 100–110 mm), in having narrower proximal subapical lamellae on fingers 3–4 (not wider than the apical lamellae versus distinctly wider), and further differs from the former species in having a tail that is not wider than the head (versus wider). Oedura nesos sp. nov. is distinguished from all nine remaining Oedura in eastern Australia by having 2 postcloacal spurs on at least one, and usually both sides, of the tail base (versus usually 1) and having a tail that is deeply depressed in cross-section (versus subcircular). Oedura nesos sp. nov. further differs from a subset of these nine eastern Australia Oedura species by having a dorsal colour pattern consisting of five distinct relatively straight edged whitish bands (versus pale v-shaped transverse bands in Oedura castelnaui Thominot, 1889, distinct darkedged ocelli or transverse blotches of varying size in Oedura coggeri Bustard, 1966, Oedura elegans Hoskin, 2019, Oedura lineata Hoskin, 2019, Oedura monilis De Vis, 1888, Oedura picta Hoskin, 2019, and Oedura tryoni De Vis, 1884, or two pale bands across the nape and base of tail in Oedura jowalbinna Hoskin & Higgie, 2008). Oedura nesos sp. nov. further differs from the remaining eastern Australian taxon Oedura argentea Hoskin, Vanderduys & Zozaya, 2018 in having a dark blackish-brown iris in life (versus silvery). Genetic differentiation. Estimates of pairwise interclade genetic distances between O. nesos sp. nov. and other members of the Oedura marmorata group from the Top End and Gulf regions range from 15–19% for mtDNA and 7–14% for UCEs, with both datasets finding the highest distances between this lineage and O. bella, and lower divergences with populations of Oedura marmorata. Phylogenetic analyses of the UCE dataset suggest that it is the sister lineage to Oedura marmorata from across the Top End region (Fig. 1A). Distribution and habitat. Only known from Groote Eylandt, off the eastern coast of the Top End region of the Northern Territory (Fig. 1B). All specimens with associated data have been collected from sandstone and it appears to be endemic to these habitats (Fig. 5). Despite significant survey effort across the island, it has not been observed or collected in open woodland habitats away from sandstone escarpment, nor observed on trees or woody debris in the vicinity of sandstone outcrops. When encountered, most geckoes were observed peering out from within crevices, potentially in ambush. Typically, shelter crevices were horizontal cracks in small boulders on bedrock. A number of geckoes were also observed foraging on open, vertical boulder faces on large, dissected sandstone pillars and outcrops. Foraging geckoes were not observed any higher than about two metres from the bedrock. Gecko species observed in sympatry include an undescribed Gehyra and Amalosia rhombifer (Gray). Suggested IUCN status. Oedura nesos sp. nov. appears to be restricted to sandstone outcrop habitats on Groote Eylandt. These habitats are only approximately 43, 919 ha (439 km 2) in extent; however, they are not under any foreseeable threat. Currently, manganese mining operations on Groote Eylandt do not pose a threat to Oedura nesos sp. nov. because manganese deposits are mostly found beneath flat, lateritic soils away from the sandstone escarpment. Where the species has been found it appears to be locally common. We therefore tentatively recommend that it be listed as Least Concern. Etymology. Nesos (Greek) meaning island, in reference to the insular distribution of this species. Used as a noun in apposition.

Oliver, Paul M., Günther, Rainer, Richards, Stephen J. (2019): Systematics of New Guinea treefrogs (Litoria: Pelodryadidae) with erectile rostral spikes: an extended description of Litoria pronimia and a new species from the Foja Mountains. Zootaxa 4604 (2): 335-348, DOI: https://doi.org/10.11646/zootaxa.4604.2.6

Gehyra paranana sp. nov. Bourke, Doughty, Tedeschi, Oliver & Moritz Litchfield spotted gecko (lineage nana 3) Figs. 5, 8, 9 Holotype. NTM R37057, an adult female collected from Dorat Road, Robin Falls area, Northern Territory (13.35278��S; 131.13361��E), on 21 September 2013 by P.M. Oliver, P. Skipwith and M. Hammer. Paratypes (12). Northern Territory: NTM R20246 (male), Butterfly Gorge, 6 km east of Daly River crossing (13.73333��S; 130.73333��E); NTM R37831 (field# CCM0651) and NTM R37832 (CCM0652) (females), Bullo River Station (15.65862��S; 129.65944��E); NTM R36578 and NTM R36579 (females), Bullo River Station (15.65766��S; 129.65933��E); NTM R36554 (male), Gurrandalng camp, Keep River National Park (15.87516��S; 129.051��E); NTM R37056 (male), Dorat Road, Robin Falls area (13.35278��S; 131.13361��E); NTM R37833 (CCM2881) (female), NTM R37834 (CCM2883) and NTM R37835 (CCM2885) (males), Tolmer Falls turnoff, Litchfield National Park (13.19654��S; 130.71394��E); NTM R37836 (CCM2936) (female) and WAM R177563 (CCM2937) (female), Florence Falls turnoff, Litchfield National Park (13.12641��S; 130.80463��E). Diagnosis. A Gehyra with moderately small body size (~ 50 mm, range 40���57 SVL), no flap of skin between limbs, dorsal half of rostral deeply furrowed with groove, single internasal usually present, 2 postnasals usually similar in size but if different then upper larger than lower, first supralabial slightly taller and narrower than second, 2 pairs of chin shields, first digit of manus and pes without claw, moderately flattened snout, snout straight to concave in lateral view, 7 divided subdigital lamellae on fourth toe, in adult males an average of 15 pre-cloacal pores (range 12���19) arranged in chevron pointing anteriorly. In life, moderately dark tan to brown background colour with pale and dark moderately large spots of similar size and not in contact on dorsum that tend to form transverse rows or coalescing to form short bars. Description. Body size moderately small (mean 49.7 mm, range 39.7���56.7 mm SVL; trunk length moderate (TrunkL/SVL 0.40, 0.36���0.44), body shape moderately robust and dorsoventrally flattened, moderately flattened head (HD/HL 0.42, 0.38���0.44) and snout, with short snout (SnEye/HL 0.44, 0.42���0.46), straight to slightly concave in lateral view, depression between rounded canthal ridges; neck moderately constricted. Limbs moderate length (ArmL/SVL 0.10, 0.08���0.12; LegL/SVL 0.11, 0.09���0.13); digits short; claw protruding from dorsal surface of expanded circular to oblong terminal toepad, no claws on anteriormost digit of manus and pes; 7 pairs of subdigitial lamellae on fourth toe (one individual with 6); 1���3 granular scales between proximate one or two rows of lamellae in 3 of 13 specimens examined. Nostrils rounded, directed laterally and slightly dorsally, contacted by rostral, supranasal, two postnasals of similar size but upper occasionally larger than lower, and first supralabial; supralabials and infralabials 7���9; rostral width ~1.7 x height with a deep furrow ~50% of rostral height, a fine medial groove extends 40���70% of the height of the scale from dorsal edge, surface of rostral with dimpled appearance; supranasals rectangular to round, dorsal edge straight or curved, ventral edge flat and in contact with rostral, supranasals typically separated medially by single small internarial scale (sometimes 2) along dorsal edge of rostral, postnasals of similar size, but upper occasionally larger than lower, first supralabial taller and narrower than second; mental narrow, angling inwards posterior to infralabials, terminating in triangular point, from 1/5 to 1/3 height of inner chin shields; outer chin shields smaller than inner (~1/2 length), with rounded lateral-posterior edges, smaller irregular-sized granular scales lateral to posterior half of outer chin shields; inner chin shields usually in contact or narrowly excluded to second infralabial; first scale of parainfralabial row of scales usually forming a notch on postero-ventral edge of second or third infralabial (if the latter, some granular scales frequently extend to contact second infralabial); chin scale arrangements frequently asymmetric. Scales on dorsum small, non-overlapping; scales near eyes becoming larger, scales on snout large and rounded; slightly enlarged row of scales above supralabials; scales on ventrum flat and ~6��� 8 x larger than those on dorsum, becoming granular anterior to arms; granular scales on gular region, increasing in size towards parainfralabial row and infralabials; scales on ventral surface of thighs and anterior to cloaca enlarged and flat; medial row of scales on tail greatly enlarged and much wider than long, bordered laterally by 1���2 rows of moderately enlarged scales, scales on dorsal and lateral surfaces of tail slightly enlarged and tending to be arranged in regular rows; regenerated tails with same scalation, but formed by more irregular scales. Males (n = 6) with an average of 15 (from 12���19) pre-cloacal pores forming a shallow chevron with the apex pointed anteriorly, pores penetrating scale, apex pore is generally present; 2 enlarged protruding cloacal spurs to either side of cloaca (females with only slightly enlarged analogous scales). Tail cylindrical and long, tapering to fine point. Colouration. In life, background colour of dorsal surfaces dark tan to medium-brown with reddish hues; widely scattered brownish-black dark bars or spots and pale white spots of similar diameter or thickness not in contact and often (but not always) tending to form alternating transverse rows; dark spots usually forming irregularly-shaped bars, but some individuals with roundish spots; midline somewhat lightened in many individuals; dark temporal streak posterior to eye usually present; large dark spots almost extend to eye level on crown but small pale spots or fine stippling extending to snout; supralabials light to heavily stippled with dark pigment, often with maculated appearance formed by hiatus of pigment near sutures; infralabials, mental, chin shields and scales on outer edge of jaw with dark stippling, infralabials often with maculated appearance as for supralabials; upper surfaces of limbs with small pale spots, small dark spots variably expressed, continuing to dorsal surfaces of digits; ventrum pale off-white with light stippling in centre and denser towards lateral edges; dorsal and lateral surfaces of original tails with alternating pale and dark rows of spots; regenerated tails with an admixture of dark and pale scales, some forming longitudinal streaks; ventral surface of tail as for ventrum. In preservative, any reddish hues are lost and background colour is a medium brown. Habitat. This species has been collected from sandstone rock formations in open woodland habitats, appearing to prefer large expanses of exposed sandstone such as large boulders and rock walls. It is sympatric with G. nana and G. koira (australis group) at northern localities. The former is a smaller-bodied species that is more commonly found on smaller boulders and amongst rock rubble, whereas G. koira is much larger and prefers boulders and vertical crevices. Distribution. Gehyra paranana sp. nov. occurs in the north-western Northern Territory (Fig. 1). In the east it is abundant in Litchfield National Park, approximately 100 km south of Darwin, south to Butterfly Gorge on the Douglas River. Westernmost records are from the vicinity of Bullo Station near the border of Northern Territory and Western Australia. Recently genotyped specimens from Bradshaw Field Training Area also confirm its presence in the intervening Wingate Range (unpublished data). Etymology. The specific name paranana refers to the morphological similarity of this species to G. nana, as in ���parallel to nana ���. Comparisons with other species. Gehyra paranana sp. nov. resembles G. nana closely, and the same comparisons to distinguish G. nana from other Gehyra above can largely be apply to G. paranana sp. nov. With respect to G. nana, all but one specimen of G. paranana sp. nov. possessed 7 subdigital lamellae on the fourth toe, whereas G. nana usually had 5 or 6. For body size, most G. paranana sp. nov. have SVL> 50 mm (maximum SVL is 56.7 mm), whereas most G. nana have SVL G. nana from the western Top End (Mt Bundy, Edith Falls, Hayes Creek) do attain larger body size, overlapping the range of G. paranana sp. nov. The pattern of spots is also useful, as G. paranana sp. nov. has larger, less distinct, pale spots and dark markings compared to G. nana (cf. Fig. 5), with the dark markings often forming short bars or networks, especially a dark streak behind the eye (absent in G. nana). Owing to the close morphological resemblance of the two species, confirmation of identity is possible with genetics by obtaining a tissue sample (e.g. tail tip) to generate mtDNA SNPs and consulting the diagnostic loci provided in Appendix 2., Published as part of Doughty, Paul, Bourke, Gayleen, Tedeschi, Leonardo G., Pratt, Renae C., Oliver, Paul M., Palmer, Russell A. & Moritz, Craig, 2018, Species delimitation in the Gehyra nana (Squamata: Gekkonidae) complex: cryptic and divergent morphological evolution in the Australian Monsoonal Tropics, with the description of four new species, pp. 201-244 in Zootaxa 4403 (2) on pages 215-217, DOI: 10.11646/zootaxa.4403.2.1, http://zenodo.org/record/1212095

Gehyra granulum sp. nov. Doughty, Palmer, Bourke, Tedeschi, Oliver & Moritz Kimberley granular-toed gecko (lineage nana 6) Figs. 5, 12–14 Holotype. WAM R176212 (CCM1270) (male), collected on 22 May 2013 by G. Bourke, J. Bragg, R. Laver and P. Title, Bell Gorge water hole, Western Australia (16.99333°S; 125.20489°E). Paratypes (10). WAM R158823 (male), Bigge Island (14.6019°S, 125.2039°E); WAM R168558 (BP01286) (female), Champagny Island (15.30°S, 124.25°E); WAM R168561 (BP01272) (female), Byam Martin Island (15.3833°S, 124.35°E); WAM R 171072 (BP00315) (male), Augustus Island (15.3900°S, 124.5931°E); WAM R171403 (male), Bachsten Creek bush camp (15.9897°S, 125.3294°E); WAM R171426 (female), Prince Regent River National Park (15.9810°S, 125.3731°E); WAM R176224 (CCM 1339) (male), Gibb River Road, roadside (17.1189°S, 125.1329°E); WAM R176245 (CCM 1492) (male), Gibb River Road (17.16653°S, 125.34761°E); NMV D75778 (male) and NMV D75774 (female), western end of Artesian Range (16.54834°S, 124.9961°E). Diagnosis. A Gehyra with small body size (average 40 mm, 35–46 mm SVL), no flap of skin between limbs, dorsal half of rostral deeply furrowed with groove, 1 internasal scale present or absent, 2 postnasals of similar size, first supralabial slightly taller and narrower than second, 2 pairs of chin shields, snout short and convex in lateral view, first digit of manus and pes without claw, 5 or 6 divided subdigital lamellae on fourth toe, proximal lamellae usually separated by 1–3 granular scales (10% lack granules), mode of 15 (range 10–19) pre-cloacal pores in males arranged in shallow chevron pointing anteriorly. Background colouration in life dull tan to brown with scattered small pale spots and larger dark blotches or irregular lines on dorsum, often forming short transverse rows. Description. Body size small (mean 40.1 mm, range 34.8–45.9 mm SVL), trunk length moderate (TrunkL/ SVL 0.40, 0.32–0.49), body shape slightly robust and dorsoventrally flattened, moderately flattened head (mean HD/HL 0.44, 0.29–0.54), with short snout (SnEye/HL 0.44, 0.36–0.52), convex to straight in lateral view, slight depression between curved poorly-defined canthal ridges; neck wide with only slight constriction. Limbs moderate length (ArmL/SVL 0.11, 0.08–0.14; HindL/SVL 0.12, 0.09–0.14); digits short; claw protruding from dorsal surface of expanded circular terminal toepad, no claws on anteriormost digit of manus and pes; 5 (occasionally 6) subdigital lamellae on fourth finger, usually 5 or 6 (a single individual with 7) subdigital lamellae on the fourth toe; first two proximal lamellae on digits separated by 1–3 granules, typically with 2 granules separating proximal pair and a single granule separating the next distal pair (Fig. 14). Nostrils rounded, directed laterally and dorsally, contacted by rostral, supranasal, two postnasals and first supralabial; supralabials 7–9; infralabials 7–8; rostral ~1.5 x wider than high with a deep furrow ~60% of rostral height, a fine medial groove extends 50–70% of the height of the scale from dorsal edge; supranasals rounded or rectangular, dorsal edge curved, ventral edge flat or curved and in contact with rostral, supranasals in contact in half of specimens, and the other half a single small internarial scale along dorsal edge of rostral present; 2 similarsized postnasals; mental narrow, usually angling inwards posterior to infralabials or continuing with straight sides to terminate in triangular point from 1/4 to 1/2 height of inner chin shields; outer chin shields smaller (~1/2 length) with rounded lateral-posterior edges, smaller irregular-sized granular scales lateral to posterior half of outer chin shields; inner chin shields in contact with second infralabial; first scale of parainfralabial row of scales usually forming a notch on postero-ventral edge of third (occasionally second) infralabial, with granular scales extending to contact second infralabial. Scales on dorsum small, non-overlapping; scales on crown are small, scales near the eyes are larger, scales on snout large and rounded; slightly enlarged row of scales above supralabials; scales on ventrum flat and ~6– 8 x larger than those on dorsum, becoming granular anterior to arms; granular scales on gular region, gradually increasing in size towards parainfralabial row and infralabials; scales on ventral surface of thighs and anterior to cloaca enlarged and flat; medial row of scales on tail greatly enlarged and much wider than long, bordered laterally by a row of moderately enlarged scales, scales on dorsal and lateral surfaces of tail slightly enlarged and tending to be arranged in regular rows; regenerated tails with irregular scales not arranged in rows. Males (n = 17) with a mode and mean of 15 (range 10–19) pre-cloacal pores forming a shallow chevron with the apex pointed anteriorly; pores penetrating scale; 1–2 enlarged protruding cloacal spurs to either side of cloaca (females with only slightly enlarged analogous scales); gravid females (36.1 mm and 37.9 mm SVL) with a single large egg. Tail cylindrical and moderately long, tapering to fine point. Colouration. In life, background colour of dorsal surfaces light to dark tan with reddish hues; dark brown irregularly-shaped blotches or bars tending to be transversely orientated; larger dark markings interspersed with more numerous scattered fine pale white spots not in contact; a subtle lightening of the background colour along midline; usually no dark temporal streak posterior to eye; scattered dark and smaller pale spots on head with small pale spots more than twice as numerous as dark spots; snout slightly dark; supralabials lightly to darkly stippled with maculated appearance formed by hiatus of pigment near sutures; infralabials, mental, chin shields and gulars with light to heavy dark stippling; upper surfaces of limbs with small pale and dark spots continuing to dorsal surfaces of digits; ventrum pale off-white with light or no stippling in centre and heavier stippling towards lateral edges; dorsal and lateral surfaces of original tails with alternating pale and dark rows of spots; regenerated tails with an admixture of dark and pale scales; ventral surface of tail as for ventrum. In preservative, subtle reddish hues lost with background colour a medium-brown. Habitat. This species has been collected from low rocks and rock faces in sandstone rock formations in open savannah woodlands with spinifex. Occasionally collected from trees, but only when in association with rocky outcrops. It is broadly sympatric with the larger-bodied species G. multiporosa, G. occidentalis, and G. pseudopunctata sp. nov. and with the more similarly sized G. spheniscus in the southwest Kimberley, and overlaps geographically with G. kimberleyi and true G. nana in the southern and eastern Kimberley. Distribution. Gehyra granulum sp. nov. occurs in the northwest Kimberley of Western Australia. On the mainland, as far north as Mitchell Plateau and extending along the coast through the Prince Regent River National Park, extending as far south as Wilinggin National Park, with records from the Durack and Cockburn Ranges to the east, and one record near Warmun community on the Great Northern Highway in the south-eastern Kimberley. Occurs on many off-shore islands of the western Kimberley, with specimens collected from Katers, Bigge, Coronation, Augustus, Caffarelli, Storr, Sunday, Long, Wulalum, Lachlan, Hidden, North West Molema, Byam Martin, Champagny, Bathurst, Irvine, Gibbings, King Hall and Koolan islands. Etymology. granulum (Latin) refers to the granular scales between the proximal subdigital lamellae that characterises this species. Comparisons with other species. Gehyra granulum sp. nov. is most similar to other small-bodied Gehyra from the Kimberley. It differs from G. nana, G. girloorloo and G. pluroporosa sp. nov. by usually possessing 1–3 small granules on the proximal subdigital lamellae and often lacks an internarial scale. From G. spheniscus, another small-bodied Gehyra with granules on the digits, it differs by number of granules on proximal lamellae (1–3 vs a small wedge of 5–7; Fig. 14), fewer pre-cloacal pores in males (10–19 vs 21–27) and different colouration (reddish-brown with large dark and small pale spots vs purplish-brown with similar-sized dark and pale spots).

Gehyra nana Storr, 1978 Northern spotted rock gecko (lineages nana 1, nana 2, nana 4, nana 7 & nanamulti from Moritz et al. 2018) Figs. 5���7, 14 Holotype. WAM R28214, an adult male collected from the King Edward River, Western Australia (���presumably in 14��52'S, 126��12'E ���) on 8 June 1965 by A.K. Lee. Paratypes (94). All WAM specimens, except as noted, from Western Australia (amendments from the type list of Ellis et al. [2018] that identified discrepancies between registration numbers provided in Storr���s description compared to the actual specimens based on archival records). R27750 [in error, R21359], R27751���52 [in error, R27551���51], R27757���62 [in error, R27557���62], R56639 [in error, R56839], Kalumburu; R28187, R28213, R28215 ���18, King Edward River; R40441, Augustus Is; R40469 (now MCZ R158546), R40470 (now MCZ R158547), R40471 (now MCZ R158548), R43147, R43157, Mitchell Plateau; R41391, Darcy Is; R41371 ���72, Heywood Is; R41428 ���31, Champagne Is; R41455, Bigge Is; R42704, 13 km NNE Argyle Downs; R43062���63, R43079���81, R56167, R56211, R56212, Crystal Creek; R43559 ���62, Pago; R44051���55, R44065 ���66, Sir Graham Moore Is; R44075, Anjo Point; R44088, Boongaree Is; R44110, South-west Osborne Is; R44144, St Andrew Is; R44157 ���65, Kingfisher Is; R44195 [in error, R99543], Melomys Is; R46668���72, R46706���07, R46710 [in error, R47010], R46883, R46884, R46831���32, R46837 ���38, Prince Regent River NR [NP]; R50330���32, R50400���03, R50414, R50465, R50507, R50675, R50783, R50797, R50802, R50811, Drysdale River NP; R56357, 2 km east of Mitchell River Falls. Of the 94 G. nana paratypes, 33 have been identified as specimens of other species as follows: G. occidentalis (10 specimens)���R44157���65, R99543; G. koira koira (1 specimen)���R50402; G. multiporosa (9 specimens)��� R41391, R41455, R44144, R46883���84, R46831���32 and R46837���38; G. granulum sp. nov. (13 specimens)��� R40441, R41371���72, R41428���31, R44088, R46668, R46672, R50783, R56167, R56211. Although belonging to different species, they maintain their status as paratypes of G. nana. Diagnosis. A Gehyra with small body size (mean 42 mm, range 34���55 mm SVL), no flap of skin between limbs, dorsal half of rostral deeply furrowed with groove, single internasal usually present, 2 postnasals of similar size, first supralabial taller and narrower than second, snout short and convex in lateral view, first digit of manus and pes without claw, mode of 6 (range 5���7) divided subdigital lamellae on fourth toe, mean of 15 (range 10���22) pre-cloacal pores in males arranged in a chevron pointing anteriorly. Background colouration dull light tan to brown with numerous small pale spots or short bars and large dark spots (not in contact) on dorsal surfaces that tend to form transverse rows on dorsum; no dark temporal streak behind eye. Description. Body size small (mean 42.4 mm, range 33.6���55.3 mm SVL; n.b. of 155 specimens measured, only 5 exceeded 50 mm), trunk length moderate (TrunkL/SVL 0.32���0.49), body shape moderately robust and dorsoventrally flattened, moderately flattened head (HD/HL 0.44, 0.41���0.65) with short snout (SnEye/HL 0.44, 0.36���0.51), convex in lateral view, slight depression between curved poorly-defined canthal ridges; neck wide with only slight constriction, enlarged calcium glands in some females. Limbs moderate length (ArmL/SVL 0.11, 0.08��� 0.13; LegL/SVL 0.12, 0.07���0.15); digits short; claw protruding from dorsal surface of expanded circular to oblong terminal toepad, no claws on anteriormost digit of manus and pes. Nostrils rounded, directed laterally and dorsally, contacted by rostral, supranasal, two postnasal scales and first supralabial; supralabials 7���10; infralabials 6���9; rostral ~1.6 wider than high with a deep furrow ~50% of rostral height, a fine medial groove extends 25���75% of the height of the scale from dorsal edge; supranasals rounded, dorsal edge curved, ventral edge flat and in contact with rostral, supranasals typically separated medially by single large or 2���3 smaller internarial scales along dorsal edge of rostral; 2 similar-sized postnasals. First supralabial taller and narrower than second; mental narrow, angling inwards posterior to infralabials, terminating in triangular point from 1/3 to 2/3 height of inner chin shields; outer chin shields smaller than inner (~1/2 length), smaller irregularsized granular scales lateral to posterior half of outer chin shields; inner chin shields usually in contact with second infralabial but can be narrowly or less commonly widely excluded; first scale of parainfralabial row of scales usually fitting into a notch on postero-ventral edge of third infralabial, but sometimes with second infralabial; chin scale arrangements frequently asymmetric. Scales on dorsum small, non-overlapping; scales near eyes becoming larger, scales on snout large and rounded; enlarged row of scales above supralabials; scales on ventrum flat and ~6��� 8 x larger than those on dorsum, becoming granular anterior to arms; granular scales on gular region, increasing in size towards parainfralabial row and infralabials; scales on ventral surface of thighs and anterior to cloaca enlarged and flat; medial row of scales on tail greatly enlarged and much wider than long, bordered laterally by 1���2 rows of moderately enlarged scales, scales on dorsal and lateral surfaces of tail slightly enlarged and tending to be arranged in regular rows; regenerated tails with same relative scale proportions but less regular than in original tails. Males with an average of 15 (range 10���22) pre-cloacal pores forming a shallow chevron with the apex pointed anteriorly, pores penetrating scale, apex pore is usually present; 2���3 enlarged protruding cloacal spurs to either side of cloaca (females with only slightly enlarged analogous scales). Tail cylindrical and long, tapering to fine point. Colouration. In life, background colour of dorsal surfaces light to dark tan or brown, with pinkish to reddish hues; scattered brownish-black dark spots and more numerous pale white spots not in contact and tending to form alternating transverse rows; dark spots of similar size or slightly larger than pale spots; dark spots tending to have irregular edges whereas pale spots have neatly rounded edges; occasionally the pale or dark markings form short bars; typically a lightening of the background colour along the midline; usually no dark temporal streak posterior to eye; large dark spots usually absent from head anterior to eyes but with small pale spots present or with fine pale stippling; supralabials darkly stippled with maculated appearance formed by hiatus of pigment near sutures; infralabials, mental, chin shields and scales on outer edge of jaw with light to heavy dark stippling; upper surfaces of limbs with small dark and pale spots, continuing to dorsal surfaces of digits; ventrum pale off-white with some light brown stippling in centre, heavier stippling towards lateral edges; dorsal and lateral surfaces of original tails with alternating rows of pale and dark spots; regenerated tails with an admixture of dark and pale scales; ventral surface of tail as for ventrum. In preservative, the reddish hues are lost and the background colour is medium brown. Habitat. This species has been collected from sandstone and other rock formations in open woodlands. Occurs under large flat rocks on open sandstone platforms as well as rocky creeks and gorges, including scree. Occasionally found on trees, but always near rocky outcrops. It is generally found among small boulders and loose rock rubble, but can occur on larger open rock faces in the absence of larger Gehyra species such as G. koira Horner or G. pseudopunctata sp. nov. Distribution. This species as redescribed here has the broadest distribution of any variegata group Gehyra in the AMT, ranging from Koolan Island in Western Australia through the Kimberley and Top End regions of the Northern Territory to Arnhem Land near the Gulf of Carpentaria (Fig. 1B). It is abundant in the northern (nana 4) and central (nanamulti) Kimberley, and extends from Koolan Island off the Yampi Peninsula east along the southern barrier ranges (King Leopold and Durack ranges), as far south as ~ 100 km south of Fitzroy Crossing, through Kununurra to the Victoria River-Bonaparte area (nana 1). Widespread in the Top End of the Northern Territory, represented by a western (nana 2) and eastern-Arnhem Land (nana 7) clades. South and east of the Roper River in the north-eastern Northern Territory, genetic analyses indicate that it is replaced by other lineages of the G. variegata group, including spotted varieties of G. versicolor Hutchinson, Sistrom, Donnellan & Hutchinson (Mt Isa region) and G. einasleighensis Bourke, Pratt, Vanderduys & Moritz (L. Ashman & Moritz, unpublished data). Etymology. Storr used the word nana (Latin), meaning small, to describe this small-bodied species relative to larger forms of Gehyra. Comparison with other species. This species complex differs from non-Australian Gehyra by the absence of webbing between third and fourth toes, absence of a skin fold along the posterior hindlimb, and moderate adult size (typical SVL G. nana complex can be distinguished from the G. australis group (which are largely restricted to the AMT) by smaller size (max SVL vs > 65 mm) and divided subdigital lamellae (vs at least some undivided). The distributions of G. nana species do not overlap those of the arid zone members of the G. variegata group with a similar spotted pattern such as G. punctata, G. pilbara Mitchell and G. montium Storr. However, these arid zone species can generally be distinguished by possession of a rich reddish background colour in life (Hutchinson et al. 2014) whereas G. nana is more light brown to tan. An exception is G. moritzi Hutchinson, Sistrom, Donnellan & Hutchinson, which is phenotypically very similar to G. nana but occurs in the south-central Northern Territory 1000 km away. Arboreal arid zone species such as G. purpurascens Storr, G. variegata and G. lazelli Wells & Wellington tend to be light purplish-brown with networks of dark lines, rather than well-defined spots and blotches as in G. nana. Diagnostic loci are presented in Appendix 2, and can be used to compare sequences from tail tips or other tissues for confident identification with minimum sequencing. Gehyra nana can be distinguished from the G. variegata group species that also occur in the AMT as follows: from G. xenopus Storr, G. spheniscus Doughty, Palmer, Sistrom, Bauer & Donnellan and G. granulum sp. nov. by the absence of granules between proximal lamellae (vs present), and in the case of the former species, also much smaller size (max SVL 55 mm vs 79 mm); from G. occidentalis by fewer subdigital lamellae on the fourth toe (5���7 vs 7���10) and smaller body size (max SVL 58 mm vs 76 mm); from G. multiporosa by the absence of dark lateral head streaks (vs present), and fewer pre-cloacal pores in adult males (10���22 vs 15���54). Gehyra nana can be distinguished from the southern Kimberley species, G. girloorloo, by a combination of dorsal colouration in life (background colour pale to dark tan vs pinkish-grey; diffuse vs clear markings), robust body (vs gracile) and tail tapering from base to tip (vs thin tail); from G. pseudopunctata sp. nov. by smaller body size (34���55 vs 47���58), more numerous pre-cloacal pores (10���22 vs 7���11), fewer subdigital lamellae on the fourth toe (5���7 vs 7 or 8) and smaller pale and dark spots on the dorsum; and from the southern Kimberley, Great Sand Desert and northern Pilbara species G. kimberleyi, by a moderately flattened head (HD/HL 0.29���0.54) vs. deep head (HD/HL 0.40��� 0.58) and small body size (34���55 mm vs 37���61 mm SVL for G. kimberleyi). The restricted distribution of the small-bodied G. pluraporosa sp. nov. overlaps that of G. nana (lineage nana 4) in the northern Kimberley (Fig. 1). However, G. pluraporosa sp. nov. possesses numerous pre-cloacal pores in males (25���26 vs 10���22), the upper postnasal is larger than lower postnasal (vs similar in size in G. nana other species) and has a dorsal pattern with alternating poorly-defined pale and dark blotches or bars, overlain with pale stippling (vs relatively well-defined pale and dark spots not in contact on plain reddish-brown background in G. nana). Gehyra nana most closely resembles G. paranana sp. nov., but differs in usually having 5 or 6 subdigital lamellae on the fourth toe whereas the latter usually has 7. In addition, the pale spots in G. nana tend to be smaller than the dark spots, whereas in G. paranana sp. nov. they are approximately equal in size, and the spots on G. nana also tend to be clearer than those of G. paranana sp. nov. (cf. Fig. 5, 7 vs 9)., Published as part of Doughty, Paul, Bourke, Gayleen, Tedeschi, Leonardo G., Pratt, Renae C., Oliver, Paul M., Palmer, Russell A. & Moritz, Craig, 2018, Species delimitation in the Gehyra nana (Squamata: Gekkonidae) complex: cryptic and divergent morphological evolution in the Australian Monsoonal Tropics, with the description of four new species, pp. 201-244 in Zootaxa 4403 (2) on pages 209-215, DOI: 10.11646/zootaxa.4403.2.1, http://zenodo.org/record/1212095, {"references":["Storr, G. M. (1978) Seven new gekkonid lizards from Western Australia. Records of the Western Australian Museum, 6, 337 - 352.","Ellis, R. J., Doughty, P. & Bauer, A. M. (2018) An annotated type catalogue of the geckos and pygopods (Squamata: Gekkota: Carphodactylidae, Diplodactylidae, Gekkonidae, Pygopodidae) in the collection of the Western Australian Museum. Records of the Western Australian Museum: in press.","Hutchinson, M. N., Sistrom, M. J., Donnellan, S. C. & Hutchinson, R. G. (2014) Taxonomic revision of the Australian arid zone lizards Gehyra variegata and G. montium (Squamata, Gekkonidae) with description of three new species. Zootaxa, 3814 (2), 221 - 241. https: // doi. org / 10.11646 / zootaxa. 3814.2.4"]}

Gehyra rohan sp. nov. Figs 2���5 Holotype. RBINS 2684 (Field No. PNG 14-88), adult male, near Lorengau, Manus Island, Papua New Guinea (2.0388S, 147.2371E), collected 3 December 2014 by J. R. Clegg, P. N.Taylor and M. M. T. Jocque. Paratypes. AMS R129497 ��� 129498, adult females, Lombrum, Los Negros Island, Admiralty Islands, Papua New Guinea (2.01S, 147. 42E), collected 15 December 1951 by N. C. Goddard; CAS 252881, adult female, south bank of Lorengau River, 2.5 km southwest of Lorengau, Manus Island, Papua New Guinea (2.0415S, 147.2594E), collected 30 May 2010 by R. Fisher; PNGNM 25220 (PNG 14-158), adult female, Yiringou Village, Manus Island, Papua New Guinea (2.0833S, 147.1167E), collected 9 December 2014 by J. R. Clegg, P. N. Taylor and M. M. T. Jocque); RBINS 2685 (PNG 14-157), adult male, Yiringou Village, Manus Island, Papua New Guinea (2.0833S, 147.1167E), collected 9 December 2014 by J. R. Clegg, P. N. Taylor and M. M. T. Jocque; SAMA R69881 (SJR15105), adult male, near Nae, Mussau Island, Papua New Guinea (1.524S, 149.739E), collected 18 October 2015 by K. Aplin. Diagnosis. Gehyra rohan sp. nov. is distinguished from other Gehyra species by the following suite of characters: very large size (adult SVL 130���150 mm), large head (HW/SVL 0.18���0.22, HD/SVL 0.11���0.14), prominent skinfolds on the anterior forelimbs and posterior hind limbs, weak lateral fold, heterogeneous dorsal scalation consisting of large rounded scales bordered by numerous much smaller rounded or triangular scales, massive digital discs with high number of wide undivided subdigital lamellae (finger IV 23���25, toe IV 22���26) that are not deeply notched or divided, rostral with near horizontal dorsal edge and not deeply notched, precloacal and femoral pores in a moderately long single continuous chevron of up to at least 40 pores, original tail without lateral serrations, slightly compressed and with a prominent medial row of enlarged subcaudals, and a prominent ring of orange scales around the eye in life. Description of holotype. Adult male. Habitus, large (SVL 140.3 mm) and robust (Fig. 2). Head triangular, robust (HW/HL 0.87), moderately long (HL/SVL = 0.23) and deep (HD/HL = 0.47). Snout long and robust (EN/ HL = 0.35). Rostral large, broadly rectangular with rounded corners, rostral groove descends approximately 35% of rostral height, bifurcates, and extends to almost contact both nares (Fig. 3). Supranasals ovoid but with distinct points at ventrolateral edges, separated by three much smaller asymmetrical (becoming smaller from right to left) squarish internasals in a transverse series. Nares bordered by rostral, first supralabial, one large supranasal and three or four postnasals. Supralabials large and squarish with rounded dorsal edges, total number to inflexion of mouth 14 (left) and 13 (right), and total number to midpoint of eye 11 (both sides). Supralabials bordered dorsally by 2���3 rows of enlarged scales. Infralabials large and squarish, total number to rictus of jaw 14 on both sides. Mental triangular, bordered by two infralabials and two large rounded postmentals (Fig. 3). Scales on dorsal surface of head tiny, irregular and slightly conical, becoming larger and flatter laterally and anteriorly. Superciliaries forming a brillar fold of small spiniform scales extending along the dorsal border of the orbit from anteroventral to posterodorsal corners. Pupil partially dilated, somewhat elliptical with limited crenulations. Body robust (TrK/SVL 0.50). Dorsal and lateral scales distinctly heterogeneous in both shape and size, generally consisting of irregularly arranged large rounded scales, bordered by numerous much smaller rounded or triangular scales, often forming a ���Star of David��� pattern; scales on nape much smaller than those on snout and torso. Ventral scales imbricate, arranged more regularly than those on dorsum, larger towards middle and posterior of venter, and tiny and granular on throat. Skin along ventrolateral edge of body loose and forming a weak fold along axilla-groin interval (Fig. 4). Precloacal and femoral pores (n = 40) arranged in a single recurved series terminating halfway along each femur. Hemipenal bulge present, moderately pronounced. Limbs robust and fleshy, with prominent lateral folds along anterior and posterior edge of forelimb, posterior edge of hindlimb, and less prominently on anterior edge of hindlimb (Fig. 4). Digits on both the fore- and hind limbs with prominent and expanded pads; terminal phalanges free and with well developed claws on all digits except finger I and toe I. Subdigital lamellae undivided, wide under expanded portion of disk, tapering and becoming narrower than toe proximal to the expanded disc (Fig. 3); total lamellar counts for all digits as follows (left/right): fingers I = 23/23, II = 24/25, III = 26/26, IV = 30/31, V = 27/26; toes I = 21/19, II = 24/26, III = 28/28, IV = 32/27, V = 24/22; total number of lamellae under expanded portion of the disk (left/right): fingers I = 22/21, II = 18/21, III = 19/19, IV = 21/24, V = 21/22; toes I = 21/19, II = 20/21, III = 22/23, IV = 23/23, V = 23/22. Webbing extending to base of disc on all digits, folded in preservative. Tail original, thin and short (107.8 mm in length) with blunt tip, much narrower than body at base; dorsal and lateral caudal scales granular and arranged irregularly, similar to dorsal scales on body; subcaudal scales distinctly enlarged, rounded, with a single medial row of 54 dilated (0.35���0.50 width of tail) ovoid scales extending full length of tail. Coloration in preservative. Dorsal and lateral surfaces grey with an indistinct chestnut wash and scattered darker grey maculations, generally corresponding to a single scale. Dorsal surfaces of limbs with coloration similar to dorsum of body, becoming distinctly darker distally and on posterior skinfolds of hindlimbs. Ventral surfaces of body, limbs and tail predominantly plain light tan. Paired pinkish-brown regions on anterior lateral edges of throat, and also forming two series of faint bars extending along lateral edges of venter, from posterior edge of insertion of the forelimbs to anterior edge of insertion of hindlimbs. Subdigital lamellae under expanded discs of all digits beige proximally, tending distinctly darker greyish-brown distally. Tail coloration as for body, but with more dense dark grey maculations on dorsal surfaces. Coloration in life. When initially captured: dorsal and lateral surfaces of head, body, limbs and tail dark chestnut brown mottled with patches of orange, light brown and off white, and with extensive black maculations, especially towards posterior region of dorsum and tail (Fig. 4). After capture: base dorsal color faded towards greyish but with same basic pattern (Fig. 4). Ventral surfaces of torso and limbs yellow, brightest anteriorly, undersurface of head brownish and digits white, transition between white of digits and yellow coloration of limbs relatively sharply defined, throat and torso both with regions of diffuse brown barring. Ventral surface of tail yellowish white with extensive dark-brown flecks. Scales around orbit forming a distinct orange ring, larger spiniform superciliary scales around dorsal edge bright reddish orange, grading to paler orange on smaller scales around ventral edge of eye. Details of holotype. Measurements (in mm): SVL 140.3; TL 107.8; TrK 69.8; HW 27.5; HL 31.6; HD 14.8; EN 11; EYE 6.5; IORB 12.4; POM 3.4; FA 16.9; CS 18.8. Meristic data: IN 3; SUPR 14; INFR 14; LAMF4 23; LAMT4 25; POR 40. Variation. Summary meristic values for all adults (2 males, 4 females) in the type series are as follows (mean, with the range in parentheses): SVL 138.4 (131.1���150.0); TL 107.2 (93.0���141.0); TrK 64.9 (58.0���72.6); HW 26.9 (24.5���29.8); HL 32.0 (30.2���35.0); HD 16.3 (14.8���18.80); EN 11.5 (10.9���13.2); EYE 7.6 (6.5���9.6); IORB 12.5 (11.4 ���13.7); POM 3.4 (3.0���3.8); FA 16.5 (15.5 ���17.3); CS 20.0 (17.7 ���22.7). Summary scalation information for these same 6 individuals are as follows: SUPR (to midpoint of eye) 10.5 (9���11); SUPR (rictus of mouth) 13.9 (12��� 14); INFR 12.7(11���14); LAMF4 22.9 (21���25); LAMT4 24.1 (22���26); POR 36.5 (33���40). The single male paratype SAMA R69881 (SJR15105) (Fig. 5) has a presumed developmental anomaly in which the pore-bearing and surrounding large scales have been shifted to the left, such that the right end of the pore series starts in the precloacal region, and extends almost fully along the left tibia, while the ventral scales on the right tibia are heterogeneous and do not appear to have formed properly. This specimen has two large, welldeveloped testes and in other respects appears to be a normal adult male. All adult specimens are similar to the holotype and share the key diagnostic traits including a bright orange ring extending around the eye, wide subcaudals under the original tail and prominent skinfolds on the arms and legs, although the prominence of the latter character varies with the angle of limb preservation. Smaller specimens tend to have a plainer ventral coloration and less obvious brown barring and mottling on the throat and ventrum in preservative, suggesting that this pattern is most pronounced in adult specimens. The largest specimens in the type series are all female (max SVL 150 versus maximum of 140 mm for the males), raising the possibility that the species is sexually dimorphic like several other Pacific Gehyra (Zug 2013), but given the very small number of males, more material is needed to confirm this. Based on photographs and field notes the dorsal coloration in life varies from quite dark chestnut brown to light grey, and the pattern of extensive but indistinct orange, greyish and brown mottling also varies in intensity. However much of this variation appears to be temporal, and single specimens vary extensively in appearance over short timescales (several hours) (Fig. 4). Comparisons. Only four other species of Gehyra approach the large size (adult SVL consistently> 120 mm) of Gehyra rohan sp. nov.: G. georgepottshaasti (Vanuatu), Gehyra marginata Boulenger, 1887 (Maluku), G. membranacruralis (Papua New Guinea) and G. vorax (Fiji). Based on published descriptions (Flecks et al. 2012) and field observations (RNF, SJR and Fred Kraus pers. com.) these taxa all lack a complete, bright orange ring around the eye in life, although occasional specimens of Gehyra vorax do have a yellow contour (as opposed to orange) around the dorsal edge of the eye (RNF pers. obs., for an example see page 172 in Ryan 1998). Gehyra rohan sp. nov. further differs from Gehyra marginata in having enlarged subcaudals under the original tail (versus absent), in having a chestnut brown iris (versus light green), and in having a less prominent ventrolateral dermal fringe on the body; from G. georgepottshaasti in having rounded postmentals (versus distinctly elongate) (Flecks et al. 2012); from Gehyra vorax in having a lower number of femoral pores in adult males (up to 40 versus 58���90) (Beckon 1992); and from Gehyra membranacruralis by its heterogeneous dorsal scalation consisting of large rounded scales separated by numerous much smaller rounded or triangular scales (versus large rounded scales only), and by having larger enlarged subcaudals (maximum anteroposterior length on adults> 2.5 mm versus Gehyra are consistently smaller than Gehyra rohan sp. nov. Gehyra baliola (Dum��ril, 1851) and Gehyra barea Kopstein, 1926 from New Guinea and surrounding islands are moderately large (max SVL up to at least 105 mm), but Gehyra rohan sp. nov. can be readily distinguished from these taxa by the presence of prominent popliteal folds on the forelimbs (versus not prominent), a rostral that is not or barely notched, and low number of large internasals (versus rostral deeply notched (u-shaped) with a moderate to high number of very small granular internasals). Gehyra serraticauda Skipwith & Oliver 2014 is also large (potentially up to 120 mm SVL), but can be distinguished by the same characters as the preceding two species, in addition to having well-developed tail serrations (Skipwith & Oliver 2014). The only congener recorded from Manus and Mussau is the widespread taxon Gehyra oceanica, from which Gehyra rohan sp. nov. is readily differentiated by its much larger adult size (> 100 mm versus versus absent), prominent popliteal skinfolds (versus absent), and distinctive orange ring around the eye in life (versus absent). The Gehyra insulensis / mutilata complex is also widespread in the Pacific, but Gehyra rohan sp. nov. is easily distinguished from this species complex by its much larger size (max SVL> 100 versus versus present). Distribution and ecology. Gehyra rohan sp. nov. is recorded from several localities across Manus Island. While most type material is from the east, one author (SJR) observed a very large Gehyra that is most likely this species at in lowland rainforest at Yeri River (2.001S, 146.819E) in north-western Manus (Fig. 6). Older material has also been collected from nearby Los Negros Island (see paratypes). This species has also been recorded from a single site on Mussau Island. The extent of its distribution, if any, beyond these islands remains unknown. Beckon (1992) reported a large Gehyra supposedly from Nauna Island near Manus (UPNG 5772), but noted that as it was collected from a banana box so its ultimate provenance was uncertain. Based on morphology, especially its high number of pores (62) Beckon further suggested that this animal is consistent with specimens from Fiji. Given uncertainty about provenance and morphology at this stage do not consider this a confirmed record of Gehyra rohan sp. nov. Gehyra rohan sp. nov. appears to be largely arboreal and is generally found in primary or disturbed lowland tropical rainforest (Fig. 6) on the trunks of large trees. It is also found on around human habitation in forested areas. Three of the authors (MJ, JRC, PT) found it to be reasonably common around Yiringou village in the interior of Manus, and two specimens were found on the same night on wooden beams below houses. The holotype was found running across a road in forest at night. The specimen from Mussau was found in a cave in disturbed forest close to the coast. One paratype (CAS 252881), from forest on the bluffs above the Lorengau River, that was initially ~6 meters high on the trunk of a tree, ���glided��� approximately 3 meters to an adjacent tree trunk when disturbed. Similar gliding or parachuting behavior has been observed in Gehyra mutilata (Heyer & Pongsapipatana 1970), and in numerous other genera of arboreal lizards, including many that lack obvious adaptations for gliding ( McGuire & Dudley 2011). Eytmology. Rohan is the Sohoniliu Village (Nali language) ���tok ples��� (local language) name for this gecko. The community of Sohoniliu Village requested that this name be used for the formal description of this species, and we thank them for their support of this work., Published as part of Oliver, Paul M., Clegg, Jonathan R., Fisher, Robert N., Richards, Stephen J., Taylor, Peter N. & Jocque, Merlijn M. T., 2016, A new biogeographically disjunct giant gecko (Gehyra: Gekkonidae: Reptilia) from the East Melanesian Islands, pp. 61-76 in Zootaxa 4208 (1) on pages 65-71, DOI: 10.11646/zootaxa.4208.1.3, http://zenodo.org/record/202010, {"references":["Zug, G. R. (2013) Reptiles and amphibians of the Pacific Islands: a comprehensive guide. University of California Press, Berkeley, 320 pp.","Boulenger, G. A. (1887) Remarks on Dr. A. Strauch's catalogue of geckos in the Zoological Museum of the Imperial Academy of St Petersburg. Annals and Magazine of Natural History, 5 (19), 383 - 388. http: // dx. doi. org / 10.1080 / 00222938709460262","Flecks, M., Schmitz, A., Bohme, W., Henkel, F. W. & Ineich, I. (2012) A new species of Gehyra Gray, 1834 (Squamata, Gekkonidae) from the Loyalty Islands and Vanuatu, and phylogenetic relationships in the genus Gehyra in Melanesia. Zoosystema, 34, 203 - 221. http: // dx. doi. org / 10.5252 / z 2012 n 2 a 1","Beckon, W. N. (1992) The giant Pacific geckos of the genus Gehyra: morphological variation, distribution, and biogeography. Copeia, 1992, 443 - 460. http: // dx. doi. org / 10.2307 / 1446204","Dumeril, C. & Dumeril, A. (1851) Museum d'Histoire Naturelle de Paris Catalogue methodique de la collection des reptiles. Gide et Baudry, Paris, 224 pp.","Kopstein, P. F. (1926) Reptilien von den Molukken und den benachbarten Inseln. Zoologische Mededelingen, 9, 71 - 112.","Heyer, W. R. & Pongsapipatana, S. (1970) Gliding speeds of Ptychozoon lionatum (Reptilia: Gekkonidae) and Chrysopelea ornata (Reptilia: Colubridae). Herpetologica, 26, 317 - 319.","McGuire, J. A. & Dudley, R. (2011) The biology of gliding in flying lizards (genus Draco) and their fossil and extant analogs. Integrative and Comparative Biology, 51,983 - 990. http: // dx. doi. org / 10.1093 / icb / icr 090"]}

Crenadactylus horni (Lucas & Frost, 1895) Central Uplands clawless geckos Figures 3 D, 4B, 8, 9 Ebenavia horni Lucas & Frost, 1895 [���Central Ranges��� of Oliver et al. (2010)] Holotype. NMV D7533, adult female, Camp 4 of the Horn Expedition, near Charlotte Waters, Northern Territory fide Dixon and Kluge (1964), donated to National Museum Victoria, Melbourne, Australia (NMV) by W.B. Spencer, April 1897. Stored in 70% ethanol at NMV. Diagnosis. A moderately large (to 34.8 mm SVL), robust species of Crenadactylus with wide (HW/HL 0.48��� 0.63) and deep (HD/HL 0.28���0.42) head with a long snout (SnL/HL 0.34���0.42). Rostral in full contact with nostril, enlarged internasal extending beyond supranasal, 2 slightly enlarged postmentals, dorsal scales homogeneous, smooth or feebly keeled, 6 pre-cloacal pores, innermost pore-bearing scales separated by an intervening scale, no enlarged tubercles on original tails. Ground colour tan and light brown; dorsal pattern consists of well-defined pale and dark longitudinal stripes, lateral zones pale tan with irregular dark brown stippling, occasionally forming 1 or 2 broken or weakly defined lateral lines; ventrum pale off-white with moderate to weak stippling. Description. The holotype is in very poor condition (shrivelled, blackened, front limbs broken off; Fig. 8) so the following description is based on other specimens in good condition. A moderately built, small-bodied gecko (to 34.8 mm SVL); body elongate, oblong in cross-section; head slightly wide (HeadW/HeadL 0.48���0.63), moderately long (HeadL/ SVL 0.27���0.32), slightly deep (HeadD/HeadL 0.28���0.42); in dorsal view, widest at ear openings; head distinct from neck with neck slightly constricted; canthal-loreal region slightly convex; snout moderately long (SnL/HeadL 0.34���0.43), tapering to rounded tip; eye moderately large; pupil vertical with four scallops; rostral ~3 times wider than tall, in broad to point contact with nostril, dorsal edge below supranasals concave to accommodate supranasals, rostral groove extends from posterior edge of rostral to ~50% of rostral height; supranasals oblong, ~1.5 times wider than long; single enlarged internasal circular, dorsal edge extends beyond dorsal edge of supranasals, occasionally a tiny additional internasal between enlarged supranasal and rostral; nostrils small, slightly smaller than loreals, directed laterally, in contact with rostral, supranasal, 3 postnasals and first supralabial; supralabials 7���9, antero-dorso edge of first supralabial in contact with nostril, postero-dorso edge notched by second postnasal and first loreal, other infralabials rectangular, wider than high, gradually decreasing in size, to corner of mouth; scales on crown small and rounded, ~one-half the size of dorsal scales, increasing in size anteriorly onto snout and supraocular regions; scales on snout flat and slightly enlarged, 2���3 times larger than scales on crown; 1 or 2 soft spines on posterior eyelid; mental triangular to trapezoidal in shape, widest anteriorly, not extending past ventral edge of first infralabial; 2���3 postmentals not slightly enlarged relative to other scales adjacent to infralabials; gulars small near infralabials, decreasing to granular, ~0.2 the size of scales on ventrum; infralabials 7���9, first largest and square-shaped, gradually decreasing in size posteriorly; ear opening small and round to elliptical in shape. Limbs short, distance between limbs close to half of SVL (ILL / SVL 0.37���0.50); scales on anterior surfaces of forelimbs broad and rounded, slightly imbricate, ~2 times the size of scales elsewhere on forelimb; scales on dorsal surface of hind limbs rounded and slightly raised; scales on anterior surface of thigh flat and circular, slightly larger in area than scales on hindlimb and elsewhere; fingers 5; toes 5; digits short, terminating in 2 enlarged apical plates, claw absent; fourth finger lamellae in 6 rows; fourth toe lamellae in 8 rows; apical plates much wider than digit. Scales on dorsum and lateral surfaces homogeneous and smooth and rounded (lacking defined keels) to feebly keeled, ovoid in cross-section, juxtaposed; scales slightly raised, highest point at posterior edge of scale, widest at midpoint of scale; scales on ventrum homogeneous, flat, imbricate, equal to 2 times the size of dorsal scales in area, widest at midpoint, with rounded posterior edge. Tail moderately long, maximum width 1/4 to 1/3 along tail, gradually tapering to a fine point, round in crosssection; scales on dorsal surface of tail ~2���3 times size compared to dorsum, homogeneous, flattened, imbricate, square to oblong in shape, arranged in travnserse rows; subcaudal scales flattened, ~2���3 times the size of dorsal tail scales; post-cloacal scales 1.5 times larger than typical ventral scales; single enlarged cloacal spur per side, ~3���4 times the size of surrounding scales; no visible pre-cloacal pores (except on 2 males, with six pores). Colouration and pattern. Ground colour tan to dark brown; a narrow silvery-grey vertebral stripe usually visible, 2���4 scales wide; vertebral stripe bordered by medium brown paravertebral stripes with darker brown margins; pale dorsolateral stripe 3���5 scales wide, narrowing on neck to ~6 smaller scales, continuing to posterodorsal edge of eye and through eye as pale canthal stripe; wide, dark dorso-lateral stripe up to 9 scales wide, medium brown with dark brown borders, on some individuals, dark borders are highly contrasting with central lighter zone giving the appearance of two separate stripes, continues anteriorly through eye as dark loreal stripe; pale lateral zone with poorly-defined brown lateral stripe; ventral edge of orbit dark brown; labials stippled with dark brown; ventrum pale with light to medium stippling. In one specimen (SAMA R22245), the dorsolateral stripe is narrow and poorly-defined. Distribution. Apparently isolated populations in the ranges of central Australia, with most records from the Northern Territory near Alice Springs and West MacDonnell and Finke Gorge National Parks (Fig. 2). As far north as Wilora (AMS R52216), and east to near the Dulcie Ranges National Park (NTM R14120). Also occurring in north-central South Australia near the Ngarutjara and Pukatja homelands. All recent and reliable records are confined to rocky areas, separated by less suitable habitat, indicating populations are confined to these areas. The presumed type locality of Charlotte Waters in the Northern Territory (indicated by a ���?��� in Fig. 2) is to the southeast of all other records, and the habitat is unlike that of the rest of this species��� records. The collection location is likely to be approximate based on the available landmarks that the Horn Expedition visited. Habitat. Collection records for several specimens mention they were taken from spinifex clumps or from under rocks near spinifex, with records also from a rocky gully, rock platforms or outcrops. Etymology. Named for William A. Horn, financer and early participant of the Horn Scientific Expedition to central Australia in 1894. Remarks. The subspecies C. ocellatus horni used to refer to the central ranges species plus Crenadactylus from the Pilbara, Cape Range and west coast owing to their strongly striped appearance relative to the type (sub)species, C. ocellatus. Along with C. ocellatus, C. horni is a more large-bodied species within Crendactylus. Individuals also possesses this species��� unique diagnostic character of an enlarged supranasal scale (Fig. 3 D) and rounded (not keeled) dorsal scales, otherwise, it is very similar to other striped species in the genus., Published as part of Doughty, Paul, Ellis, Ryan J. & Oliver, Paul M., 2016, Many things come in small packages: Revision of the clawless geckos (Crenadactylus: Diplodactylidae) of Australia, pp. 239-278 in Zootaxa 4168 (2) on pages 250-253, DOI: 10.11646/zootaxa.4168.2.2, http://zenodo.org/record/262140, {"references":["Lucas, A. H. S. & Frost, C. (1895) Preliminary notice of certain new species of lizards from central Australia. Proceedings of the Royal Society of Victoria, 7, 264 - 269.","Oliver, P., Adams, M. & Doughty, P. (2010) Molecular evidence for ten species and Oligo-Miocene vicariance within a nominal Australian gecko species (Crenadactylus ocellatus, Diplodactylidae). BMC Evolutionary Biology, 10, 386. http: // dx. doi. org / 10.1186 / 1471 - 2148 - 10 - 386","Dixon, J. R. & Kluge, A. G. (1964) A new gekkonid genus from Australia. Copeia, 1964, 174 - 180. http: // dx. doi. org / 10.2307 / 1440848"]}

Oedura Gray, 1842 Type species: O. marmorata Gray, 1842, by monotypy. Generic diagnosis. A genus of the Diplodactylidae distinguished from all related genera by the possession of enlarged juxtaposed dorsal scales approximately the same size as the ventrals (versus much smaller in related genera). Further distinguished from other taxa formerly placed in Oedura (Amalosia, Hesperoedura and Nebulifera; Oliver et al. 2012), by the combination of large size (adult SVL>60 mm), usually more than one cloacal spur per side, and dorsal pattern generally including a weak to bold series of transverse bands or disjunct blotches and no evidence of a vertebral stripe., Published as part of Oliver, Paul M. & Doughty, Paul, 2016, Systematic revision of the marbled velvet geckos (Oedura marmorata species complex, Diplodactylidae) from the Australian arid and semi-arid zones, pp. 151-176 in Zootaxa 4088 (2) on page 162, DOI: 10.11646/zootaxa.4088.2.1, http://zenodo.org/record/258925

Crenadactylus Dixon & Kluge, 1964 Type species. Diplodactylus ocellatus (= Crenadactylus ocellatus) Gray, 1845 by original designation. Diagnosis. Based on Dixon & Kluge (1964). Species within the genus Crenadactylus are small (max SVL 35 mm) geckos differing from all other gekkonids by a combination of the following external morphological characteristics; digits with enlarged subdigital lamellae, terminal phalangeal elements forked and claws absent from all digits; outer margins of anterior portion of frontal notched to receive posterior projection of paired nasals. Internal morphological diagnostics include palatines short and broad, atlas fused dorsally; stapes imperforate (stapedial foramen absent); two pairs of sternal ribs, one or two pairs of mesosternal ribs; 28 sacral and presacral vertebrate; sacral diapophyses overlapping and fused (not fused in juveniles); in adults anterior tip of mesoscapula fused to precoracoid process at its union with precoracoid; interclavicle dagger-shaped; fingers 5; toes 5; phalangeal formula of manus 2-3-4-5-3, pes 2-3-4-5-4; 24���26 scleral ossicles; 13 or 15 premaxillary teeth; 29���31 maxillary teeth; 33���37 mandibular teeth; cloacal bones present in males (Dixon & Kluge 1964). Etymology. A combination of the Latin word crena meaning ���notch��� and Greek word daktylos meaning ���finger��� in reference to the forked terminal phalanges. Content. Crenadactylus horni (Lucas & Frost, 1895), Crenadactylus naso Storr, 1978, Crenadactylus occidentalis sp. nov., Crenadactylus ocellatus (Gray, 1845), Crenadactylus pilbarensis sp. nov., Crenadactylus rostralis Storr, 1978, Crenadactylus tuberculatus sp. nov., Published as part of Doughty, Paul, Ellis, Ryan J. & Oliver, Paul M., 2016, Many things come in small packages: Revision of the clawless geckos (Crenadactylus: Diplodactylidae) of Australia, pp. 239-278 in Zootaxa 4168 (2) on page 247, DOI: 10.11646/zootaxa.4168.2.2, http://zenodo.org/record/262140, {"references":["Dixon, J. R. & Kluge, A. G. (1964) A new gekkonid genus from Australia. Copeia, 1964, 174 - 180. http: // dx. doi. org / 10.2307 / 1440848","Gray, J. E. (1845) Catalogue of the Specimens of Lizards in the Collection of the British Museum. Trustees if the British Museum / Edward Newman, London, xxviii + 289 pp.","Lucas, A. H. S. & Frost, C. (1895) Preliminary notice of certain new species of lizards from central Australia. Proceedings of the Royal Society of Victoria, 7, 264 - 269.","Storr, G. M. (1978) Seven new gekkonid lizards from Western Australia. Records of the Western Australian Museum, 8, 337 - 352."]}

Oedura bella sp. nov. Gulf marbled velvet gecko Figs. 5, 7, 8 Holotype. QM J94016 (field number���OMAR #001), adult male with original tail, 10 km south of Mt Isa on Boulia Road, Queensland, Australia (20.8617��S, 139.4617��E), collected by P.M. Oliver, M. Vucko and M. Vickers, on 20 February 2007. Paratypes. Northern Territory: AMS R53437���8, 37 km north of McArthur River base camp on Borroloola Road (16.10��S, 136.12��E); AMS R53467, Caranbirini Water Hole, 21 km north of McArthur River base camp (16.22��S, 136.15��E); AMS R53643, Glyde River, 10 km east of McArthur River base camp (16.43��S, 136.17��E); AMS R53782, 37 km north of McArthur River Camp (16.10��S, 136.12��E); SAMA R34188, McArthur River Station (16.67��S, 135.85��E). Queensland: NTM R21288, Musselbrook Reservoir (18.61��S, 138.08��E); SAMA R34208���9, SAMA R35425, Lawn Hill NP (18.58��S, 138.50��E); QM J52748, Lawn Hill Station, Century Project Site (18.75��S, 138.58��E); QM J74927, Hells Gate (17.47��S; 138.37��E); QM J75207���8, Lawn Hill (18.71��S, 138.48��E). Referred material. AMS R138727���8, Groote Eylandt (13.83��S, 136.42��E); NTM R7494���5, Umbakumba Rd (13.88��S, 136.50��E); NTM R7541, Ayakamindadina (13.97��S, 136.60��E). Diagnosis. A medium-sized (SVL: mean 78 mm, max 92 mm) species in the O. marmorata complex with a wide (HW/SVL 0.19���0.23) and moderately deep head (HD/SVL 0.10���0.12), short body (Trk/SVL 0.41���0.49), short original tail (TL/SVL 0.49���0.65) that is narrower than head and slightly depressed, rostral usually less than half divided, terminal lamellae moderately wide (ToeW/SVL 0.021���0.030), proximal subdigital lamellae of all fingers not wider than apical pair, 12���17 precloacal pores in males and base colouration dark purplish brown with 5 distinct to faint light dorsal bands from nape to hindlimbs. Description. A medium-sized (to 92.0 mm SVL) and moderately robust Oedura (Trk/SVL 0.41���0.49). Head moderately narrow (HW/SVL 0.19���0.23) and not strongly depressed (HD/SVL 0.10���0.12). Rostral 25���50% divided, bordered dorsally by two internasals, internasals bordered dorsally by two oblong moderately enlarged supranasals and 1���3 (mode 1) smaller intervening scales. Enlarged supralabials 9���10 to midpoint of eye, 11���13 in total; infralabials 9���13. Limbs moderately long (ArmL/SVL 0.12���0.15; LegL/SVL 0.14���0.16). Subdigital lamellae moderately expanded and prominent, 7���8 rows (mode 8) under finger three and 7���9 toe three, distal lamellae on all digits either divided or deeply notched, apical distinctly larger and generally wider than more proximal pairsespecially on fingers, proximal lamellae undivided. Enlarged postcloacal spurs 2 or 3, in a single cluster per side. Original tail relatively short (TL/SVL 0.49���0.65) and wide (TW/TL 0.18���0.28), gradually tapering to a point, slightly depressed in cross-section and slightly concave ventrally; moderately wide but shows considerable variation depending on body condition (TW/SVL 0.10���0.15), scalation relatively homogeneous with dorsal and ventral scales of similar size. Fully regrown tails shorter (TL/SVL 0.44���0.53) and wider (TW/TD 0.21���0.31) than original tails. In preservative, dorsal colouration of adults variable; base colouration dark purplish brown, with 5 distinct to very indistinct broad light bands from nape to hindlimbs, with a further 4���7 similar bands on the tail. Light bands with little to extensive dark brown flecking or blotches; on larger specimens dark brown pigmentation is extensive such that the broad light bands are reduced to two narrow indistinct and often broken bands. The anterior (nuchal) light band extends across the nape and joins or approaches a light lateral band extending from the labial scales and above the tympanum. The canthal stripe ranges from distinct to indistinct. Dark brown regions between light bands and elsewhere on the dorsal and lateral surfaces of the head, torso, limbs and tail, with extensive further light flecking and blotches. Venter relatively plain light buff with a very faint wash of light brown, particularly towards the lateral extremities and on the head and limbs, more extensive brown mottling also present along the ventrolateral edges of the tail. Regrown tails dark brown with varying amounts of light flecking, but no clear pattern. Hatchlings and small juveniles lack light flecking and generally have a very simple pattern of five sharplydefined light bands on the neck and body, and 5���7 light bands on the tail, separated by broad plain dark brown regions. As specimens grow, the sharp definition of light and dark regions decreases, and the amount of light flecking and blotching across the body increases. In life, the base colouration of adults is dark purplish brown during the day, washed out grey at night; light bands and flecks ranges tend to be bright yellow. The canthal stripe tends to be very strongly defined and a white labial stripe is also sometimes clearly apparent. Iris very dark brown. Particulars of holotype. QM J94016 (field tag number���OMAR#001), male with original tail (in mm): SVL 77.0, HW 14.6, HD 7.4, HL 19.6, EN 6.3, IN 2.8, IO 6.7, EYE 5.0, TrK 31.0, ArmL 10.1, LegL 10.9, TL 38.0, TW 8.3, TD 5.6, 3FW 1.8, 3TW 2.3. Scales: RC 25%, SuL 12(10), InF 11, CS, 3FL 7, 3TL 8, precloacal pores 16. Distribution and habitat. Restricted to the ranges around the south and western edges of the Gulf of Carpentaria; from the Selwyn Range around Mt Isa in the southeast, north to Riversleigh and Musselbrook regions in the Queensland-Northern Territory border, as far west as the McArthur River and Borroloola regions of Northern Territory, with a further apparently isolated insular population on Groote Eylandt to the north (see comments), and another isolated record from the northern edge of the Barkly Tablelands (Fig. 2). Field observations suggest this species is primarily saxicoline, using both horizontal screes and vertical faces (including road cuttings), however, it has also been recorded under bark around the base of trees (but in rocky country) (G. Bourke, pers. comm.). Etymology. From the Latin masculine adjective bellum (used in its feminine form), meaning amongst other things pretty, handsome, charming, fine, lovely, neat, pleasant, agreeable, active, gallant or good. In reference to the very attractive contrasting yellow and dark colour pattern of this species. Comparisons. Oedura bella is similar to parapatric populations of O. marmorata in northern Australia (although it differs in both nuclear and mitochondrial loci). It differs in external morphology by possessing an original or regrown tail that is much narrower (TW/SVL 0.10���0.15 versus 0.19���0.24) and generally less than the width of head, and O. bella also reaches a smaller maximum size, although adult sizes overlap (adult SVL usually 77���92 mm versus 77���97 mm) and sample sizes are low. Oedura bella differs from O. cincta and Western (see below) by the combination of its smaller maximum size (SVL 64���92 mm versus 77���106 mm), shorter original tail (TL/SVL 0.49���0.65 versus 0.58���0.80), and its narrower terminal lamellae (0.21���0.30 versus 0.23���0.36) and narrower lamellae series on the fingers (not wider than terminal lamellae versus wider on digits 3 and 4). It further differs from the geographically proximate Oedura cincta by generally having a rostral partially divided by a crease (versus usually fully divided). Oedura bella can be distinguished from the three species of Oedura in the Kimberley region in Western Australia by having subdigital lamellae that are slightly expanded around the midpoint of the digit (versus strongly tapering in O. gracilis King, 1984, or obviously flared and often as wider or wider than the apical lamellae in O. filicipoda King, 1984 and O. murrumanu Oliver, Laver, Melville & Doughty, 2014), and its moderately long and slightly swollen tail (versus very long [approaching length of body] and tapering in gracilis, or greatly flattened and wider than body in O. filicipoda and O. murrumanu). With a maximum SVL of 92 mm, O. bella is also smaller than O. filicipoda and O. murrumanu (which both regularly exceed 100 mm). Oedura bella differs from all other Oedura in eastern Australia by possessing more than one postcloacal tubercle and in having a base colouration of five relatively thin dorsal bands. The latter character distinguishes it from two other small saxicoline Oedura in eastern Australia: O. coggeri has large ocelli on limbs and torso and O. jowalbinna has a pale pinkish gray dorsum with distinct dark-edged bands across the neck and base of tail and a plain yellow original tail (Hoskin & Higgie 2008). All other Oedura in eastern Australia tend to be larger (SVL> 90 mm) and also have dorsal patterns that do not feature thin light bands; specifically, O. castelnaui has wide bands, O. monilis has blotches or ocelli, and O. tryoni has dense small spots. Remarks. No genetic samples were available from the Groote Eylandt samples that we refer to this species. Given their geographic disjunction from all other material, the distinctiveness of this population warrants further investigation., Published as part of Oliver, Paul M. & Doughty, Paul, 2016, Systematic revision of the marbled velvet geckos (Oedura marmorata species complex, Diplodactylidae) from the Australian arid and semi-arid zones, pp. 151-176 in Zootaxa 4088 (2) on pages 165-168, DOI: 10.11646/zootaxa.4088.2.1, http://zenodo.org/record/258925

Couper, Patrick J., Oliver, Paul M. (2016): A new species of gecko from arid inland regions of eastern Australia (Diplodactylus; Diplodactylidae). Zootaxa 4093 (4): 525-538, DOI: http://doi.org/10.11646/zootaxa.4093.4.4

Crenadactylus occidentalis sp. nov. Western clawless geckos Figs. 4 C, 10, 11 [���Carnarvon Basin��� of Oliver et al. (2010)] Holotype. WAM R113683, adult male, Dirk Hartog Homestead, Dirk Hartog Island, WA (26��00'S, 113��12'E), collected by B. Maryan and R. Browne-Cooper, 19 April 1992. Fixed in 10% formalin, stored in 70% ethanol at WAM. Paratypes (6). WAM R57525, 40 km north-east of Yuna, WA (28��06'S, 115��15'E); WAM R96676, 10 km north-west of Wandina Homestead, WA (27��56'S, 115��33'E); WAM R 120779, Nerren Nerren Station, WA (27��03'24"S, 114��35'21"E); WAM R124891, 38 km west south-west Hamelin Homestead, WA (26��35'34"S, 113��53'22"E); WAM R131376, 70 km south of Exmouth, WA (22��34'47"S, 114��07'00"E); WAM R135497, False Entrance Well, WA (26��23'S, 113��19'E). Diagnosis. A medium-sized (to 32.6 mm SVL) species of Crenadactylus with wide (HW/HL 0.50���0.60) head. Rostral in full contact with nostril, internasal (if present) not extending beyond supranasal, 2 (occasionally 3) small postmentals, dorsal scales homogeneous and weakly keeled, 3���4 pre-cloacal pores, innermost pore-bearing scales separated by an intervening scale, no enlarged tubercles on original tails. Ground colour tan and light brown; dorsal pattern consists of well-defined pale and dark longitudinal stripes, small pale white to orange spots comprised of 2��� 3 scales may be present, lateral zones pale to dark grey with at most faint uniform stippling (not forming lines); ventrum pale off-white with stippling absent to moderate. Description of holotype. WAM R113683, a small adult male with the following meristics: SVL 28.0 mm; tail absent, remaining portion ~ 3.1 mm; HeadL 8.1 mm; HeadW 4.6 mm; HeadD 2.7 mm; SnL 2.5 mm; ILL 12.6 mm; SupLab 9 (left), 8 (right); InfLab 9, 8; internasals 0; PCP 4. A slender, elongate, small-bodied gecko; body oblong in cross section, ~2.0���2.5 times wider than deep; head narrow (HeadW/HeadL 0.57), elongate (HeadL/ SVL 0.29), moderately depressed (HeadD/HeadL 0.33); in dorsal view, widest at ear opening, as wide as widest portion of body; head not distinct from neck, neck weakly constricted; loreal region weakly convex, concave around nasal region; snout elongate (SnL 2.5; SnL/HeadL 0.31), rounded tip; eye moderately large; rostral ~3.0 times wider than high, in broad contact with nostril, dorsal edge concave to accommodate first supranasal, slightly notched by median internasal, rostral groove originates from dorsal edge of scale, ~40% of rostral height; internasals absent; supranasals 2 per side, first oblong, transverselyoriented, ~2.5 times wider than high, in broad contact with nostril; second supranasal small, rounded, ~half the size of first supranasal; nostril small, slightly smaller than second supranasal, directed dorsolaterally, in contact with rostral, first and second supranasals, three postnasals and first supralabial; postnasals 3; supralabials 9 (left), 8 (right), antero-dorso edge of first supralabial in contact with nostril; second supralabial square, narrower than first; supralabials 3���4 square, equal in size, 5���9 rectangular, gradually decreasing in size, all wider than high; scales on crown small and granular, increasing in size anteriorly onto snout; scales on snout flat, enlarged; supraocular scales enlarged; two enlarged conical scales at postero-dorso edge of eye; mental trapezoid-shaped, widest anteriorly, narrowing slightly to posterior edge, posterior edge not extending beyond ventral edge of first infralabials; postmentals 2, round in shape, equal to size of scales posterior to postmentals; gulars rounded, gradually decreasing in size posteriorly to granular scales; infralabials 8 on each side, 1���3 largest, equal in size, square-shaped, slightly higher than wide, 4���8 rectangular, wider than high, gradually decreasing in size posteriorly. Limbs short, distance between limbs nearly half of SVL (ILL / SVL 0.45); scales on dorsal surface of forelimbs imbricate, rounded and slightly raised, becoming flatter distally; scales on ventral surface of forelimbs smaller; scales on dorsal surface of hind limbs imbricate, rounded and slightly raised, becoming flatter distally; scales on ventral surface of hind limbs similar to scales on dorsal surface of hind limbs, becoming granular distally; fingers 5 (fifth finger missing from left forelimb); toes 5; claws absent from all digits, digits moderately long; fourth finger lamellae in 8 rows; fourth toe lamellae in 10 rows; in dorsal view, enlarged scale between apical plates ~2 times the size of adjacent lateral scales, apical plates much wider than digit; two pairs of dorsal apical plates; one pair of ventral apical plates, ovate. Scales on dorsum small and homogeneous, round to oblong, juxtaposed; scales weakly keeled, highest point at posterior edge of scale, widest at midpoint of scale, becoming flatter and imbricate laterally; scales on ventrum homogeneous, flat, imbricate, ~2���3 times the size of dorsal scales in area, widest at midpoint, rounded posteriorly. Tail detached, remaining portion ~ 3.1 mm, tapers gradually from widest point posterior to vent; round in crosssection; scales similar to scales on dorsum, rounded and slightly raised, imbricate; ventral tail scales flattened and imbricate, ~1.5���2.0 times the size of scales on dorsal surface of tail; single enlarged raised post-cloacal spur on both sides of cloaca, ~10 times the size of surrounding scales; pre-cloacal pores 4, 2 innermost pore-bearing scales separated by a single intervening poreless scale, pore in notch on posterior edge of scale. Colouration and pattern. In preservative, dorsum with longitudinal series of stripes, yellowish-brown ground colour; vertebral stripe silvery-grey, 3 scales wide; 2 dark brown paravertebral stripes, 3 scales wide; pale wide dorsal stripes, light yellow-brown with scattered dark scales, 4 scales wide, terminates at postero-dorsal edge of eye; brown latero-dorsal stripes 2���3 scales wide, originates at posterior edge of eye, over ear and above limbs, continuing to base of tail. Variation. Most specimens were similar to the holotype in body size and proportions and scalation. Variation in the snout tip area usually took the form of from 3���6 scales in contact with first supranasals, including presence or absence of a small internasal between the first supranasals (Fig. 3 C). Pre-cloacal pores were 4 with the exception of WAM R 120789 with 3. Males possessed enlarged cloacal spurs whereas in females, the spurs are smaller and protrude away from body less. Original tails with large scales (~4���6 times the size of dorsal scales), scales rectangular with rounded posterior edge, slightly imbricate, arranged in regular rows; subcaudal scales ~1.5 times larger than dorsal tail scales, flattened, imbricate; regenerated tails with smaller, flatter scales compared to original scales; hemipenes bifid. Colouration and pattern. Highly contrasting dark brown and pale yellowish-brown longitudinal stripes; ground colour a dull tan or light yellow-brown; vertebral stripe pale silvery-grey, 3 scales wide, extending to crown; vertebral stripe enclosed by dark brown paravertebral stripes, 3 scales wide, that meet between hindlimbs and base of tail, extending anteriorly to crown, becoming intermixed with pale scales on crown; wide pale tan to light yellow-brown dorsolateral stripe, 4 scales wide, extends from upper part of eye to tail, extending forwards through eye to form pale canthal stripe, edged with dark brown in some individuals; above canthal stripes a dark brown triangular patch on snout; rostral and labial scales pale, darkly stippled to varying degrees, posterior edge of orbit with dark brown mark; dark brown dorsolateral stripes, 2���3 scales wide, emanate from posterior edge of eye and continue to tail, extending forwards through eye to form dark brown loreal stripe; pale dorsolateral zone occasionally flecked with small orange clusters of scales; dark paravertebral and dorsolateral stripes peppered with fine clusters or single pale scales in some individuals; lateral zone pale silvery-white, almost continuous to ventrum, thin weakly-defined brown lateral stripe usually present, below lateral stripe, scattered dark flecks may be present. Ventrum ranges from an almost immaculate, pale silvery-grey, to lightly stippled with black, occasionally forming short streaks. Limbs variably marked, a mixture of pale, light brown and dark brown scales with scattered clusters or ocelli of 3���8 pale scales, darker scales forming irregular dark variegations. Original tails have a continuation of the dorsal pattern, with dark brown paravertebral stripes on dorsum merging to form single dark vertebral stripe; regenerated tails have a mixture of dark and pale scales, not forming a regular pattern. Distribution. Occurs on the Carnarvon coast of Western Australia from areas inland of Geraldton in the south, extending north through Kalbarri, Shark Bay and to the southern part of the North West Cape at Yardie Creek (where it is narrowly sympatric with C. tuberculatus sp. nov.) (Fig. 2). Also collected from Bernier and Dorre islands in Shark Bay and off Carnarvon. Furthest inland occurrences near Wandana Nature Reserve at the southern part of its distribution (~ 100 km inland) and Nerren Nerren further north (~ 70 km inland) (both with genotyped specimens), otherwise most records occur within 50 km of the coastline. Habitat. Collection records indicate a preference for spinifex and coastal dune vegetation such as shrubs, Acacia and Banksia. Occasionally collected from under tin and limestone slabs. Etymology. The specific name occidentalis (Latin) refers to the western distribution of this species. Remarks. This species��� distribution is closely associated with the western coast and its furthest inland extent is also the most southern, possibly indicating a low tolerance for extremely dry conditions. This area receives variable rainfall during the winter months (May���August), with decreasing rain furthest north where C. occidentalis is only found in a narrow band along the coast. The sister species of C. occidentalis is C. tuberculatus sp. nov., with an estimate of divergence around 5 mya (Oliver et al. 2010). Unlike the rather typical Crenadactylus morphology possessed by C. occidentalis, however, C. tuberculatus sp. nov. has a highly derived morphology and pattern and is entirely restricted to the rugged limestone hills and gorges of the Cape Range. The two species come into contact at the northern edge of the distribution of C. occidentalis, including sympatry where the two species meet at Yardie Creek., Published as part of Doughty, Paul, Ellis, Ryan J. & Oliver, Paul M., 2016, Many things come in small packages: Revision of the clawless geckos (Crenadactylus: Diplodactylidae) of Australia, pp. 239-278 in Zootaxa 4168 (2) on pages 253-256, DOI: 10.11646/zootaxa.4168.2.2, http://zenodo.org/record/262140, {"references":["Oliver, P., Adams, M. & Doughty, P. (2010) Molecular evidence for ten species and Oligo-Miocene vicariance within a nominal Australian gecko species (Crenadactylus ocellatus, Diplodactylidae). BMC Evolutionary Biology, 10, 386. http: // dx. doi. org / 10.1186 / 1471 - 2148 - 10 - 386"]}

Doughty, Paul, Ellis, Ryan J., Oliver, Paul M. (2016): Many things come in small packages: Revision of the clawless geckos (Crenadactylus: Diplodactylidae) of Australia. Zootaxa 4168 (2): 239-278, DOI: http://doi.org/10.11646/zootaxa.4168.2.2

Oedura fimbria sp. nov. Western marbled velvet gecko Figs. 5, 9, 10 Holotype. WAM R154783, adult male with original tail, Brockman Ridge, Western Australia, 23.3108��S, 119.9169��E, Western Australia, collected by J. Fraser on 5 April 2004. Paratypes. All from Western Australia: WAM R105965, 7 km north of Mount Magnet (28.00��S, 117.88��E); WAM R119991, Hope Downs (23.01��S, 119.10��E); WAM R119993, Hope Downs (23.00��S, 119.12��E); WAM R129595, 120 km north-west of Newman (22.92��S, 118.88��E); WAM R129635, 120 km north-west of Newman (22.92��S, 119.02��E); WAM R135369, Mount Brockman (22.31��S, 117.32��E); WAM R154796���7, Walga Rock (27.40��S, 117.47��E); WAM R157504, WAM R157508, WAM R157516, Packsaddle Range (22.92��S, 118.89��E); WAM R157595, West Angelas (23.19��S, 118.86��E); WAM R160074, 32.5 km east-south-east of Meentheena Outcamp (21.33��S, 120.750��E); WAM R165150, 5 km north-north-west Python Pool (21.32��S, 117.23��E). Referred material. All from Western Australia: WAM R52852, 12 km east of Tallering Peak (28.10��S, 115.75E); WAM R84004, 6 km north of Mount Magnet (28.03��S, 117.85��E); WAM R84365, WAM R84366, 17 km north-north-east Anketell homestead (27.90��S, 118.95��E); WAM R87544, 30 km south-south-west Glenburgh homestead (24.68��S, 115.00��E); WAM R97012, Woolgerong Rock (27.40��S, 117.38��E); WAM R106289, 7 km north Mount Magnet (28.00��S, 117.88��E); WAM R119086, Virgin Springs, Carnarvon Range (25.10��S, 120.72��E); WAM R119837, Yandicoogina (22.72��S, 119.02��E); WAM R129622, 120 km north-west Newman (22.92��S, 118.88��E); WAM R132296, Ulongunna Rock (27.12��S, 117.23��E); WAM R132626, Burrup Peninsula (20.60��S, 116.81��E); WAM R135445, Mount Brockman (22.31��S, 117.32��E); WAM R146593, WAM R146594, 40 km south-east Pouyouwuncubban (22.15��S, 119.02��E); WAM R157508, Packsaddle Range (22.92��S, 118.89��E); WAM R160066 58 km east-south-east Meentheema Outcamp (21.32��S, 121.00��E); WAM R165150, 5 km northnorth-west Python Pool (21.32��S, 117.23��E). Diagnosis. A large (mean SVL 94 mm, max 104 mm) species in the O. marmorata complex, with a moderately broad head (HW/SVL 0.18���0.21), moderately long body length (Trk/SVL 0.42���0.53), tail moderately long (TL/SVL 0.65���0.80), narrower than head and roughly circular in cross-section, rostral 25���60% divided, apical lamellae wide (TW/SVL 0.027���0.036), subdigital lamellae in a flared series wider than apical pair on fingers 3���4, 13���26 precloacal pores in adult males, and basic dorsal colouration (especially on juveniles) usually including 5 light transverse bands. Description. A large (to 104 mm SVL) and moderately elongate Oedura (Trk/SVL 0.40���0.53); head moderately wide (HW/SVL 0.18���0.21) and deep (HD/SVL 0.09���0.13). Rostral 25 to 60% divided, bordered dorsally by two nasals, nasals bordered dorsally by two relatively small supranasals and 2���4 (mode 3) small intervening scales. Supralabials 9���11 to midpoint of eye, 11���13 in total; infralabials 10���13. Limbs moderate (ArmL/SVL 0.12���0.15, LegL/SVL 0.14���0.17). Subdigital lamellae expanded and prominent, 7���10 under third finger, 8���10 under third toe, distal lamellae on digits either divided or deeply notched, apical pair distinctly larger (but not always wider) than and separated from other pairs, penultimate pair slightly narrower, subsequent pairs on digits 2���5 distinctly flared, proximal lamellae undivided. Original tail moderately long (TL/SVL 0.65���0.80) and narrow (TW/TL 0.13���0.17) with a slight ventral groove, varying from somewhat depressed to almost circular in cross-section, relative width and depth varies greatly depending on body condition; caudal scalation homogeneous. Fully regrown tails shorter (TL/SVL 0.51���0.66) and wider (TW/TL 0.18���0.26) than original tails. In preservative, base colouration of dorsum dark purplish brown, with a highly variable amount of light cream flecking or blotching. Most adult specimens have 5���6 distinct to very indistinct pale light transverse dorsal bands with wide brown central regions, with a further 5���8 similar bands on the tail. In larger specimens, however, all traces of bands have been lost. The anterior (nuchal) light band generally joins or approaches a light lateral stripe that extends from the labial scales and above the tympanum. Bands are typically very faded and indistinct on adult specimens from the north of the range, but remain more conspicuous in specimens of all sizes from the south. Extensive further light flecking and blotches are present between the bands, and elsewhere on the dorsal and lateral surfaces of the head, torso and limbs; northern specimens tend to have many smaller flecks, while southern specimens have fewer larger flecks or reticulate blotches. On a small number of apparently aberrant specimens, the light pigmentation is very extensive and brown areas form narrow transverse bands. Venter plain light buff, sometimes with faint brownish tinge on the throat and the terminal lamellae. Regrown tails dark brown with varying amounts of light flecking, but no clear bands. Hatchlings and small juveniles are dark purplish brown dorsally, with 5 sharply-defined light cream transverse bands on the torso, 5���7 additional light bands on the tail, and a narrow white labial stripe. The dorsum of the head tends to be lighter brown than the remainder of the body. With increasing size, the light transverse bands become less clearly defined and usually develop a wide brown mid-region. An additional very light creamish transverse band or series of blotches also develops at the approximate midpoint of the pre-existing light bands; light flecking and blotching across the body, limbs and tail also becomes increasingly apparent. In life, the basic pattern and colouration of adults matches those of preserved specimens, however, the darker regions are purplish during the day, silvery grey at night, and light regions tend to be whitish to relatively bright yellow. Iris very dark brown. Particulars of the holotype. WAM R154783, adult male with original tail (in mm): SVL 104.0, HW 19.3, HD 10.9, HL 24.6, EN 8.1, IN 3.7, IO 8.5, EYE 5.4, TrK 49.0, ArmL 13.2, LegL 16.4, TL 74, TW 12.2, TD 10.2, 3FW 3.1, 3TW 3.6. Scale counts: SuL 10(12), InF 10, CS 2/3, 3FL 9, 3TL 8, precloacal pores 26. Hemipenes everted, right 10.2 mm long, left 7.6 mm long, pores in an almost continuous curved series separated by 2 poreless medial scales, two additional pore-bearing scales are also present posterior to the main series in the precloacal region. Distribution and habitat. Occurs in the Pilbara, Gascoyne and Murchison regions of mid-western Western Australia (Fig. 2). In the north (Pilbara), this species extends to the southern edge of the Great Sandy Desert, east to Karlamilyi National Park, and west to the Burrup Peninsula. In the south it occurs in the Gascoyne, western Murchison and Yalgoo regions, east as far as the Barlee Range, south to Mt Magnet and Gullwa, and east to the Little Sandy Desert. From collector���s notes associated with voucher specimens, this species was recorded as occurring on relatively massive ranges, rocky outcrops and breakaways, caves and gorges 114 times, whereas there are only 4 occurrences on trees and one under tin. We have only observed this species on rocky outcrops with large boulders (Pilbara) or on cliff faces (Kennedy Range). Etymology. Fimbria is Latin for fringe in reference to the lateral fringes of expanded lamellae along the sides of the digits. Used as a noun in apposition. Comparisons. Oedura fimbria sp. nov. is most similar to O. cincta and shares moderately large size (SVL> 100 mm), moderately long tail which tends towards rounded in cross-section and is not wider than the head, wide flared subdigital lamellae, dorsal pattern usually consisting of numerous poorly-defined light flecks and blotches and often thin light bands. The only clear diagnostic morphological character between the two speices is an incomplete rostral crease on O. fimbria sp. nov. (25���60% versus 60 [rarely]���100% [usually] of the rostral height). There are some differences in average for relative head length and body length, but ranges overlap extensively (see Table 1). Oedura fimbria sp. nov. can be distinguished from other lineages in the O. marmorata species complex from northern Australia by possessing a longer (TL/SVL 0.65���0.80 versus 0.53���0.63) and narrower tail (always narrower than the head). It differs from O. bella in its larger size (max SVL 104 mm versus 92 mm), longer tail (TL/SVL 0.65���0.80 versus 0.49���0.65), wider terminal lamellae (3TW/SVL 0.23���0.36 versus 0.21���0.30) and prominently flared subdigital lamellae series���especially on fingers 3 and 4 (proximal lamellae wider than apical lamellae versus not wider). Oedura fimbria sp. nov. can be distinguished from the three other species of Oedura in the Kimberley region in Western Australia as follows: from O. gracilis by its moderately long and swollen tail (versus very long [approaching length of body] and tapering) and in having subdigital lamellae series that is flared around the midpoint of the digit (versus tapering); from O. filicipoda in having a narrow tail that is not wider than the head and near circular in cross-section (versus wider and very flattened) and smaller eyes (EYE/SVL 0.051���0.064 versus 0.071���0.076); and from O. murrumanu in having a smaller eye (EYE/SVL 0.051���0.064 versus 0.067���0.069) and a longer rostral crease (> 25% [typically 40���60%] versus, Published as part of Oliver, Paul M. & Doughty, Paul, 2016, Systematic revision of the marbled velvet geckos (Oedura marmorata species complex, Diplodactylidae) from the Australian arid and semi-arid zones, pp. 151-176 in Zootaxa 4088 (2) on pages 168-171, DOI: 10.11646/zootaxa.4088.2.1, http://zenodo.org/record/258925

Gehyra kimberleyi B��rner & Sch��ttler, 1983 Robust termitaria gecko Figs. 2���4 Holotype. BSRC Geck 65 SC (private collection of Achim-R��diger B��rner, Germany), ��� 5 km south of Derby, W.A.��� Diagnosis. Digits broadly expanded basally and subdigital scansors present on all digits of manus and pes. Digit I of manus and pes clawless, penultimate phalynx of digits II���V free from scansorial pad. Differs from non- Australian Gehyra by the combination of: absence of webbing between third and fourth toes, absence of a skin fold along the posterior hindlimb and medium adult size (SVL Gehyra by the combination of moderate size (SVL Details of holotype. From original description (in mm). SVL 36; TailL 39; HeadL (tip of rostral to anterior corner of ear) 9; HW 7.3; HD 4.3; HD/SVL 0.119; SupLab* 6 / 7; InfLab* 6 / 7; FingSubDL 6; ToeSubDL 9; rostral with cleft, undivided; first SupLab higher than second (*not defined, possibly measured to centre of eye). Description of species. Range of variation for measurements and scale counts are presented in Table 1. A medium-sized (adult SVL mean 50.4 mm, range 42.0��� 61.3 mm), moderately robust gecko. Head deep (HD/HL 0.48, 0.40���0.58), moderately wide (HW/HL 0.86, 0.76���0.97), widest posterior to eye, tapering to a rounded snout, narrowing posteriorly to a slightly constricted neck. Snout short (SnEye/HL 0.42, 0.39���0.46), generally shorter than twice eye length (OrbL/SnEye 0.58, 0.54���0.63), occasionally slightly concave on dorsal surface, canthus rounded; in profile view, snout convex with blunt tip. Body moderately long (TrunkL/SVL 0.42, 0.37���0.47) and robust, slightly depressed. G. kimberleyi G. girloorloo sp. nov. G. nana (south Kimberley) G. pilbara N = 19 N = 17 N = 40 N = 14 (M = 11 /F = 8) (M = 10 /F = 7) (M = 20 /F = 20) (M = 4 /F = 10) On head, dorsal scales largest on snout, ~ 2.5 times larger than scales on crown of head and body and slightly projecting and imbricate. Nostrils large and circular, bordered by rostral, supranasal, two postnasals of similar size and first supralabial; supranasals round to triangular in shape, usually separated by a single large internarial scale. Rostral scale wider than high ( RosH/RosW 0.64, 0.56���0.74), slightly gabled, usually with a deep medial cleft ~ 60 % of rostral height. Supralabials 7���9 (mode 8), infralabials 7���8 (mode 7). First supralabial higher than subsequent supralabials. Supralabials bordered dorsally by enlarged row of scales posteriorly that decrease in size as for supralabials. Mental scale roughly triangular, divides postmentals 20���40 %. Postmentals in contact with first infralabial only; single pair of outer chin shields smaller with rounded outer edge, ~two-thirds the height of postmentals, in contact with first and second infralabials, thereby excluding postmental from contact with second infralabial. Second or third infralabial notched and in contact with first parinfralabial. Eye small (OrbL/HL 0.24, 0.21���0.27), circular, small fringe of mostly rounded scales extends from the postero-dorsal to the anterio-ventral edge. Ear openings moderately large and circular. Dorsal scales homogeneous, small, rounded and juxtaposed; gular scales granular; ventral midbody scales 2���3 as large as dorsal scales, slightly imbricate and tending to be diamond-shaped; limbs covered in granular scales, scales on anterior surfaces enlarged, especially on legs. Pre-cloacal pores 12���17, mode 16, perpendicular to body distally and medial curving forwards to form a sharply pointed ���v��� with anteriorly-directed apex. Usually a single small post-cloacal spur present on each side in males. Females lack pre-cloacal pores and prominent post-cloacal spurs. Limbs very short (ArmL/SVL 0.11, 0.09���0.13; HindL/SVL 0.12, 0.10���0.17); five digits on each limb, claws present on digits 2���5, claws long and free, arising from dorsal surface of expanded toe pads and extending above and well beyond toe pad, row of elongate and sharply pointed distal scales on periphery of dorsal surface of toe; toe pads ovoid; subdigital lamellae divided, 5���7 (mode 6) pairs on fourth finger, and 6���9 (mode 7) on fourth toe, apical lamellae wedge-shaped. Tail moderately long and moderately thin to slightly swollen, slightly dorso-ventrally flattened in crosssection, tapering to fine tip. Dorsal scales of tail flattened with rounded posterior edge, tending to be arranged in rows. Enlarged, wide series of scales beneath tail, bordered peripherally by 2���4 rows of enlarged scales, decreasing in size on lateral surfaces. Colouration in preservative. Background dorsal colour varies from dark reddish- or greyish-brown with fine dark stippling to light salmon-pink, pale and patternless in larger individuals (Fig. 2). Dorsal surfaces of head, torso and tail usually with scattered poorly-defined small, dark greyish-brown spots, blotches, flecks or short bars; pale grey spots interspersed among the darker markings, usually forming rows of 4���6 spots; on the neck and crown, blotches are usually separated, on the torso often transversely aligned to form loosely-defined bands on the body and on the tail often forming concentric bands. Dark longitudinal streaks on lateral surface of neck posterior to eye or a series of small dark blotches. Limbs with similar pattern to dorsum, but generally less well defined. Ventral colour pale, lightly stippled but more darkly pigmented anteriorly, especially in gular region. Iris pale silvery blue, pupil vertical and pale with crenelated edge. Original tails with alternating dark and pale concentric rings or series of blotches. Regenerated tails with no clear pattern, although indistinct longitudinal brown streaks are sometimes apparent. Colouration in life. Based on photographs of specimens captured in the south-western Kimberley. The dorsal background colour is pale or golden tan to medium dark brown, and the dorsal pattern consists of scattered dark brown blotches and smaller pale spots that occur in alternating rows, especially the vertebral zone, with smaller spots occurring on the head, flanks and limbs. The dorsal pattern tends to be less distinct in larger specimens (Fig. 3). Habitat and reproduction. Habitat notes from collectors of specimens held at the WAM indicate most were collected from termite mounds (128 of 227 such records). Other collection locations indicate generalist habits, with specimens collected from trees, under logs and other ground cover and including vertical rock faces. Like other species in the Gehyra variegata-punctata species-group (Doughty 1996), G. kimberleyi lays a single egg. Distribution. Occurs throughout the southern Kimberley region, from Cape Leveque at the northern edge of the Dampier Peninsula in the west, as far north as Mt Elizabeth in the central Kimberley, and north-east to Lake Argyle near Kununurra (Fig. 1). Most records are from south of the King Leopold and Durack barrier ranges, with several records south of the Fitzroy Crossing area. There are no records north of the Durack Range in the eastern Kimberley, although this region is the most poorly surveyed of the Kimberley. In the Northern Territory, the species is recorded from Buchanan Station. There are records of genotyped specimens as far south as the northern Pilbara (15 km north-east of Nifty Mine and Meentheena Nature Reserve), with several specimens from along the 80 mile beach extension of Dampierland between the west Kimberley and Pilbara regions. In the Great Sandy Desert, there are records from Dragon Tree Soak Nature Reserve in the south to records further north. The Great Sandy Desert remains poorly surveyed and this species may be more widely distributed in this region. Etymology. Although the original description mentions the ���Kimberleys��� (sic) (B��rner & Sch��ttler 1983, p. 1), no explicit etymology was provided. The presumed correct formation of the specific name is ���kimberleyensis���, which means the taxon is from the Kimberley region. The proposed name ���kimberleyi��� would refer to a male named Kimberley. The error, however, cannot be emended under current ICZN rules of nomenclature. The suggested common name refers to species��� body habitus, and ���termitaria��� indicates this species��� habitat preference (although it is not exclusive to termitaria). Comparison with other species. Gehyra kimberleyi can be distinguished from all non-Australian Gehyra by the combination of absence of webbing between the third and fourth toes (versus present), the absence of a skin fold along the posterior hindlimb (versus present) and its smaller size (max SVL versus > 65 mm). Within Australia, G. kimberleyi can be distinguished from all members of the G. australis species-group (which are largely restricted to the AMT) by its smaller size (max SVL versus max SVL> 65 mm), divided subdigital lamellae (versus at least some undivided) and single egg per clutch (versus two). Gehyra kimberleyi can be distinguished from most other members of the G. variegata-punctata group that also occur in the AMT as follows; from G. xenopus Storr and G. spheniscus by the absence of a wedge of granules between proximal lamellae (versus present), and in the case of the former species, smaller size (max SVL 62 versus 79 mm); from G. occidentalis Storr by its smaller body size (max SVL 62 versus 76 mm), deeper head, postmentals 3 x longer than wide (versus 4 x), lower number of subdigital lamellae (6���8 versus 7���10), fewer pores in males (12���17 versus 18���30) and more clearly-defined spots on the dorsum; and from G. multiporosa Doughty, Palmer, Sistrom, Bauer & Donnellan, 2012 by fewer pores in adult males (12���17 versus 20���49), dark markings on dorsum not tending to form transverse bars and the absence of dark lateral head streaks. Gehyra kimberleyi can be distinguished from both geographically proximate and type G. nana by short snout and deep head, tan or greyish brown dorsal colouration (versus reddish), generally larger body size (SVL 42.0��� 61.3 versus 35.0��� 44.3), more larger, irregular and pale blotches (versus small and clearly defined), diffuse and transversely oriented dark brown blotches on dorsum (versus clearly defined brown blotches or spots). Gehyra kimberleyi overlaps with a number of other members of the G. variegata-punctata species-group from the Australian arid zone, especially in the northern Pilbara. It differs from species in this group in having dorsal markings of dark blotches that tend to coalesce to form transverse bars (versus continuous longitudinal lines or network in G. variegata and G. montium ��� Hutchinson et al. 2014), and by having 7���9 supralabial scales (versus 8���10 in G. variegata and 8���11 in G. montium ��� Hutchinson et al. 2014). It is distinguished from members of the Gehyra punctata species complex by its diffuse dorsal colouration (versus well defined dark and light spots or blotches) and postmentals that do not contact the second supralabial. From Gehyra pilbara (with which it has frequently been confused) it can be distinguished by its larger size (max SVL 62.3 versus 45.1 mm), tall, narrow postmentals (versus extremely short and wide; cf. Fig. 4), and tan or greyish-brown dorsal colouration (vs. reddish-brown)., Published as part of Oliver, Paul M., Bourke, Gayleen, Pratt, Renae C., Doughty, Paul & Moritz, Craig, 2016, Systematics of small Gehyra (Squamata: Gekkonidae) of the southern Kimberley, Western Australia: redescription of G. kimberleyi B��rner & Sch��ttler, 1983 and description of a new restricted range species, pp. 49-64 in Zootaxa 4107 (1) on pages 51-55, DOI: 10.11646/zootaxa.4107.1.2, http://zenodo.org/record/260932, {"references":["Borner, A. R. & Schuttler, B. I. (1983) An additional note on the Australian geckos of the genus Gehyra. Miscellaneous Articles in Saurology, 12, 1 - 4.","Doughty, P. (1996) Allometry of reproduction in two species of gekkonid lizards (Gehyra): effects of body size miniaturization on clutch and egg sizes. Journal of Zoology, 260, 703 - 715. http: // dx. doi. org / 10.1111 / j. 1469 - 7998.1996. tb 05316. x","Doughty, P., Palmer, R., Sistrom, M. J., Bauer, A. M. & Donnellan, S. C. (2012) Two new species of Gehyra (Squamata: Gekkonidae) geckos from the north-west Kimberley region of Western Australia. Records of the Western Australian Museum, 27, 117 - 134.","Hutchinson, M. N., Sistrom, M. J., Donnellan, S. C. & Hutchinson, R. G. (2014) Taxonomic revision of the Australian arid zone lizards Gehyra variegata and G. montium (Squamata, Gekkonidae) with description of three new species. Zootaxa, 3814 (2), 221 - 241. http: // dx. doi. org / 10.11646 / zootaxa. 3814.2.4"]}

Crenadactylus rostralis Storr, 1978 South-west Kimberley clawless geckos Figs. 3 H, 4F, 16, 17 Crenadactylus ocellatus rostralis Storr, 1978 [���Kimberley A��� of Oliver et al. (2010, 2012b)] Holotype. WAM R32154, adult male, Geikie Gorge, Western Australia (18��07'S, 125��39'E), collected by W.H. Butler, April 1968. Fixed in 10% formalin, stored in 70% ethanol at WAM. Paratypes (17). WAM R23058 *, 43 km SE Halls Creek, WA (18��27'S, 127��15'E); WAM R26824���26, Napier Range, WA (17��20'S, 124��50'E); WAM R27388 ���92, Mount Anderson, WA (18��02'S, 123��56'E); WAM R46058 ��� 59*, near Wolfe Creek Meteorite Crater, 122 km SW Halls Creek, WA (18��43'S, 126��45'E); WAM R46115 *, 11 km E Margaret River Homestead, WA (18��38'S, 126��58'E); WAM R56841 ���45*, Lake Argyle, WA (16��15'S, 128��45'E). Some registration numbers presented in the original description are in error, with WAM R56441���45 associated with different species. The error originates from a minor numerical error in the registration numbers presented by Storr (1978), and underlined here. The correct numbers associated with the paratype specimens are WAM R56841��� 45. Nine paratypes (marked with an asterisk) designated by Storr (1978) fall within the distribution of lineages associated with the C. naso complex identified by Oliver et al. (2012b), and are not true C. rostralis. Diagnosis. A moderate-sized (to 32.7 mm SVL) species of Crenadactylus. Rostral not in contact with nostril, 2���3 internasals present, not extending beyond supranasal, postmentals granular, dorsal scales homogeneous and weakly keeled, 4���6 pre-cloacal pores, innermost pore-bearing scales in contact, no enlarged tubercles on original tails. Ground colour light greyish brown; dorsal pattern consists of well-defined pale and dark longitudinal stripes, lateral zone light grey with usually two moderate to well-defined dark lines between limbs, stippled with single pale scales; ventrum pale grey to white with moderate stippling forming longitudinal lines. Description of holotype. WAM R32154, adult male with the following meristics (in mm): SVL 29.8; TailL 27.5 (original); HeadL 9.3; HeadW 4.6; HeadD 2.8; SnL 3.3; ILL 14.5; SupLab 9 (left), 9 (right); InfLab 9, 9; internasals 3. A slender, small-bodied gecko; body elongate, oblong in cross section, ~1.5���2 times wider than deep; head narrow (HeadW/HeadL 0.50), elongate (HeadL/ SVL 0.31), moderately depressed (HeadD/HeadL 0.30); in dorsal view, widest between eye and ear openings, narrower than widest portion of body; head not distinct from neck, neck weakly constricted; loreal region weakly convex, weakly concave around nasal region; snout elongate (SnL 3.3; SnL/HeadL 0.36), rounded tip; eye moderately large; rostral ~3 times wider than high, excluded from nostril contact by first supranasal, dorsal edge concave to accommodate first supranasal, slightly notched by median internasal, rostral groove originates from dorsal edge of scale below internasal, ~40% of rostral height; internasals 3, median internasal tiny; supranasals 2 per side; first oblong, transversely-oriented, ~2.0���2.5 times wider than high, in broad contact with nostril; second supranasal small, rounded, ~size of first supranasal; nostril small, ~1/3 size of first supranasal, directed dorso-laterally, in contact with both supranasals, 3 postnasals and first supralabial; postnasals 3, first and third ~half the size of first supranasal, second small, ~1/3 of first and third; supralabials 9 on both sides, antero-dorso edge of first supralabial in contact with nostril; second supralabial rectangular, wider than high, supralabials 3���5 smaller than first and second, all wider than high, sixth supralabial ~equal to first supralabial, supralabials 7���9 smaller than first two, gradually decreasing in size posteriorly; scales on crown small and granular, ~half the size of dorsal scales, becoming flattened and increasing in size anteriorly onto snout; scales on snout flat, enlarged, ~1.5���4.0 times the size of scales on crown; supraocular scales enlarged; 3 enlarged conical scales at postero-dorso edge of eye; mental trapezoidal, widest anteriorly, narrowing slightly to rounded posterior edge; gulars granular, ~1/3 to 1/4 the size of scales on ventrum; infralabials 9 on each side, first trapezoidal, slightly smaller than second, infralabials 2���4 square, gradually decreasing in size posteriorly, infralabials 5���9 rectangular, wider than high, gradually decreasing in height and width posteriorly. Limbs short, distance between limbs nearly half of SVL (ILL / SVL 0.48), just over half of tail length (ILL / TailL 0.52); scales on dorsal surface of forelimbs flat and rounded, imbricate, decreasing in size and becoming more granular distally; scales on ventral surface of forelimbs similar to dorsal; scales on dorsal surface of hind limbs rounded and slightly raised, imbricate, decreasing in size and becoming more granular distally; scales on ventral surface of hind limbs circular and slightly raised, increasing in size and becoming flatter and imbricate distally, granular on feet; fingers 5; toes 5; claws absent from all digits; digits moderately short; fourth finger lamellae 6 rows; fourth toe lamellae 8 rows; in dorsal view, enlarged scale between apical plates ~equal to 2 times the size of adjacent lateral scales; apical plates much wider than digit; two pairs of dorsal apical plates; one pair of ventral apical plates, circular. Scales on dorsum homogeneous, small, feebly keeled, juxtaposed, scales slightly raised with highest point at posterior edge of scale, widest at midpoint of scale, becoming flatter and imbricate laterally; scales on ventrum homogeneous, flat, imbricate, equal to 1.5���2.0 times the size of dorsal scales in area, widest at midpoint, rounded posteriorly. Tail original, nearly equal to SVL (TailL/ SVL 0.92), tapers gradually from widest point posterior to vent to narrow rounded point; round in cross-section; scales homogeneous, imbricate and flat, variable in size and shape, ~3���4 times the size of scales on dorsum; ventral tail scales similar to dorsal tail scales, flattened and imbricate, ~1.5���3 times the size dorsal tail scales; single enlarged spur on both sides of cloaca, ~5���6 times the size of surrounding scales; pre-cloacal pores 6, forming a slight curve, median innermost pore-bearing scales in contact, pore-bearing scales notched on posterior edge to accommodate pore. Coloration and pattern. In preservative, colour faded to light brown. A thin pale vertebral stripe 2���3 scales wide, bracketed by medium brown paravertebral stripes 3���4 scales wide; a light brown dorsolateral zone 6���8 scales wide, with several widely-spaced pale scales, continuing forwards through eye and onto snout as pale canthal stripe; dark brown dorsolateral stripes 3���4 scales wide, in contact with dorsal edge of hindlimb, continuing forwards to eye and onto snout as loreal stripe; light brown lateral zone below dark dorsolateral stripe, 3���4 scales wide; ventro-lateral stripe medium brown, weakly defined, 3���5 scales wide; crown and snout medium brown, bordered by light brown stripe (extension of dorsolateral zone); supralabials stippled with dark brown, continuing as weakly-defined dark brown stripe to arm; ventrum pale with 4 longitudinal stripes, outer pair darker, 2���3 scales wide, inner pair narrower, 1���2 scales wide; arms with dark variegations on dorsal surface, legs with weaklydefined dark brown longitudinal lines, ventral surfaces of limbs with irregular stippling; basal portion of tail (original) with dark brown stripes continuing from paravertebral and dorsolateral stripes on body, ventral portion of tail with 2 dark brown stripes. Variation. Other specimens generally corresponded to the holotype specimen, with the following exceptions. The degree of keeling on the dorsal scales ranged from feebly keeled to smooth and rounded. Females lacked pre-cloacal pores and the spurs were smaller and less projecting. Colouration and pattern. In life (based on recent photographs by PMO), background colour grey-brown with a series of alternating pale and dark alternating longitudinal stripes; pale grey vertebral stripe bordered by medium dark brown paravertebral stripes; pale to medium grey dorsolateral zone, continuing forwards through eye as pale canthal stripe; dark brown lateral stripe, continuing through eye on snout as poorly-defined loreal stripe; light grey ventrolateral zone with occasionally poorly-defined thin dark stripe near ventrolateral edge of body; top of head dark brown with some pale mottling; labials pale, stippled with dark brown. Dark brown paravertebral and lateral stripes usually with scattered pale scales. Lateral stripe usually darker than paravertebral stripes, but can also be of similar shade. Ventrolateral zone can be much paler than dorsolateral zone. Limbs dark brown with variable small pale spots; original tails continue dorsal pattern, regenerated tails with a mixture of pale and dark scales not forming conspicuous lines. In preservative, pattern consists of alternating dark and light stripes. Silvery-grey vertebral stripe, narrow, 2���3 scales wide; medium grey-brown paravertebral stripes, bordered by dark brown edges, 6���7 scales wide; pale dorsolateral stripe, 4���5 scales wide, starts at postero-dorsal edge of eye, continues above limbs to tail; dark brown dorso-lateral stripe extends posteriorly from back of eye to top of hindlimb, anteriorly extends to top of ear opening; below dorsolateral line are two thin, ventro-lateral stripes; pale lateral zone divided by thin dark lateral stripe between limbs, forming pale line below dorsolateral that extends from angle of jaw across top of forelimbs and through hindlimb. Ventrum lightly stippled, one or two weakly defined brown lines towards outer edge of ventrum. Pale canthal and dark loreal stripes weakly defined, continuation of pale and dark dorsolateral stripes; top of eyes purplish; labials pale and heavily stippled with brown flecks; posterior edge of jaw heavily stippled, continuing as lateral line below ear opening to forelimb. Limbs dark, back of thigh has strong pale stripe, diffuse with small clusters of pale scales. Distribution. Genotyped specimens were all collected from the Devonian Reefs Ranges in the south-western Kimberley region (Fig. 2), extending from the Napier Ranges in the north-west to the Pillara Range on Gogo Station in the south-east. Additional (not genotyped) records of Crenadactylus from Mt Anderson in the south-west (~ 40 km from Derby) and from the south-east Kimberley are here considered to belong to C. rostralis pending genetic data to resolve their status (Fig. 2). All samples formerly regarded as this species from elsewhere in the Kimberley are here considered part of the C. naso species complex. Habitat. The few records with collection details all mention occurring in spinifex, including a number of recent samples that were raked from large clumps on and around limestone ranges (PMO, pers. obs.). Etymology. The name rostralis is from the Latin rostrum, meaning beak or snout, in reference to this species��� distinctive configuration of having the rostral scale separated from the nostril., Published as part of Doughty, Paul, Ellis, Ryan J. & Oliver, Paul M., 2016, Many things come in small packages: Revision of the clawless geckos (Crenadactylus: Diplodactylidae) of Australia, pp. 239-278 in Zootaxa 4168 (2) on pages 262-264, DOI: 10.11646/zootaxa.4168.2.2, http://zenodo.org/record/262140, {"references":["Storr, G. M. (1978) Seven new gekkonid lizards from Western Australia. Records of the Western Australian Museum, 8, 337 - 352.","Oliver, P., Adams, M. & Doughty, P. (2010) Molecular evidence for ten species and Oligo-Miocene vicariance within a nominal Australian gecko species (Crenadactylus ocellatus, Diplodactylidae). BMC Evolutionary Biology, 10, 386. http: // dx. doi. org / 10.1186 / 1471 - 2148 - 10 - 386","Oliver, P. M., Doughty, P. & Palmer, R. (2012 b) Hidden biodiversity in rare northern Australian vertebrates: the case of the clawless gecko (Crenadactylus, Diplodactylidae) of the Kimberley. Wildlife Research, 39, 429 - 435. http: // dx. doi. org / 10.1071 / WR 12024"]}

Diplodactylus ameyi sp. nov. Eastern Deserts Fat-tailed Gecko Figs: 2, 3, 4 A, 5 A, 6, 7 (top row) [Diplodactylus] ��� conspicillatus ��� F (Oliver et al. 2009) Holotype. QM J 90778 (Fig. 2), 3.4km NNE Noonbah homestead (24 ��04' 51 "S, 143 �� 11 ��� 54 ���E) Queensland, Australia, frozen tissue sample at South Australian Museum ABTC 113844, collected by Mark Hutchinson. Paratypes. Queensland. SAMA R 63336, Winton, (22 �� 27 'S, 142 �� 57 'E); QM J 92287, Winton (22 �� 28 ' 42 "S, 142 �� 53 ' 31 "E); AMS R 110564, Camp 14km NE Scott���s Tank, Diamantina Lakes, NW of Windorah (23 �� 45 'S, 141 �� 40 'E); AMS R 110529, Scott���s Tank, Diamantina Lakes, NW of Windorah (23 �� 58 'S, 141 �� 32 'E); QM J 83467, Noonbah Station (24 ��07'S, 143 �� 11 'E); QM J 90774, Noonbah homestead, 3.4km NNE (24 ��04' 51 "S, 143 �� 11 ' 54 "E); QM J 90170, Valetta Stn (24 �� 15 ' 21 "S, 143 ��05' 57 "E); QM J 59978 Jundah. ���Noonbah Stn��� Rd to 'Waterloo Stn' in Bore Paddock (24 ��07' 40 "S, 143 �� 11 ' 30 "E); QM J 56888, Waterloo site 1 (24 �� 16 ' S, 143 �� 13 'E); AMS R 143856, Stonehenge area, within 10km N to S of Stonehenge (24 �� 22 'S, 143 �� 19 'E); QM J 89191, Tyrone, approx 70km NW of Charleville - 3km S of old north Tyrone homestead (25 �� 58 ' 55 "S, 145 �� 44 ' 17 "E); QM J 35697, Ambathala, 1km S Ra Tank (26 ��01' 30 "S, 145 ��04' 30 "E); QM J 79909, Mariala (26 ��05'S, 145 ��04'E); QM J 74874, Mariala Nature Ref. (26 ��05' 30 "S, 145 ��04' 15 "E). New South Wales. AMS R 158426, Sturt NP, Silver City Hwy, Wittabrinna Ck. Crossing (29 �� 22 ' 38 "S, 142 ��02'08" E); AMS R 132996, AMS R 132997, Wanaaring, 4km W of Wanaaring at Turnoff to Wilcannia (29 �� 42 'S, 144 ��07'E); AMS R 141988, Wanaaring (29 �� 42 'S, 144 ��09'E); AMS R 165698, Nocoleche Nature Reserve, 11km W of Wanaaring - Wilcannia Rd (29 �� 52 '08"S, 144 ��00' 34 "E); AMS R 165659, AMS R 165697, AMS R 166837, Nocoleche Nature Reserve, 11km West of Wanaaring - Wilcannia Road (29 �� 52 '08"S, 144 ��00' 34 " E); AMS R 162733, AMS R 165673, Lake Peery National Park (30 �� 43 ' 28 "S, 143 �� 29 ' 15 "E). ������continued on the next page Species-group diagnosis. A member of the Diplodactylus conspicillatus group as defined by the following combination of characters: most supralabials small and granular, at most only one enlarged anterior (first) supralabial; terminal lamellae on fingers at most only slightly wider than digit; prominent enlarged subdigital lamellae absent; tail short, as wide or wider than body, depressed with heterogeneous scalation, and bearing large plate-like scales and/or conical tubercules arranged in transverse rows; and dorsal colouration extremely variable, but lacking large well defined bands or blotches. Specific diagnosis. A large member of the D. conspicillatus group (max SVL = ~ 60 mm) in which the first supralabial is small and does not contact the ventral edge of nasal scale (Fig. 3). Snout broad and ���U���-shaped snout (dorsal view, Fig. 4 A), convex along its dorsal edge and lacking a well-defined canthus rostralis (Fig. 5 A). The canthal stripe is absent or poorly defined (Figs 1, 2, 4 A, 5 A, 6 & 7 - top row). Description. SVL mm 46.0��� 60.2 (n = 22, mean = 53.2, SD = 4.3). Proportions as % SVL: AG 42.2���55.9 (n = 22, mean = 48.9, SD = 3.4); T 33.2���44.7 (n = 15, mean = 37.8, SD = 3.0). Head: broad and deep but not strongly differentiated from neck; snout longer than diameter of eye, broad and blunt (U-shaped when viewed from above; Fig. 4 A), with a conspicuous bulbous swelling anterior to the eyes (resulting in a poorly defined canthus rostralis), and gently convex along dorsal edge in lateral profile (Fig 5 A). HW 76.9���92.3 % HL (n = 22, mean = 83.8, SD = 4.5); HD 44.8 ���57.0% HL (n = 22, mean = 48.9, SD = 3.2); S 42.5���48.6 % HL (n = 22, mean = 45.4, SD = 1.15); EE 24.0��� 32.1 % HL (n = 22, mean = 28.2, SD = 2.4); covered in small granular scales; rostral shield large and lacking a medial groove, hexagonal with 8���13 scales contacting posterior edge (n = 22, mean = 10.5, SD = 1.3); mental shield hemispherical; 10���15 scales contacting posterior edge (n = 22, mean = 12.1, SD = 1.3); supralabial scales 11���18 (n = 22, mean = 15.0, SD = 2.0), the first not enlarged and subequal with the rest of the supralabial row which are not differentiated from the adjacent loreal scales (Fig. 3); infralabial scales 12���19 (n = 22, mean = 15.1, SD = 1.9), small and barely differentiated from adjacent chin scales; eye large, pupil vertical with crenulated margin; ear small, round to horizontally elliptic. Neck: broad with small granular scales on dorsal surface that are only slightly larger than the adjacent scales on the lateral surfaces. Trunk: moderate and somewhat stout; scales of dorsum plate-like and markedly larger than the granules on flanks; scales granular on ventral surface but increase in size on pectoral region; precloacal pores absent; a small cluster of postcloacal tubercles present in both sexes but larger and more prominent in males. Limbs: moderate; forelimb 28.3���37.1 % SVL (n = 22, mean = 32.5, SD = 2.5); hindlimb 31.1���40.8 % SVL (n = 22, mean = 35.5, SD = 2.6); digits moderate with no or only slight distal expansion; subdigital scales granular with a small pair of apical lamellae; 9���13 scales beneath fourth finger (n = 22, mean = 11, SD = 1.0); 9��� 14 scales beneath fourth toe (n = 22, mean = 11.7, SD = 1.19). Original tail: short 33��� 45 % SVL, width 48.0��� 78.8 % of tail length (n = 15, mean = 61.1, SD = 7.9); spade-like and bluntly pointed (lacking an acute attenuated tip); dorsal scales each with a short blunt to sharp medial tubercle, arranged in clear transverse series which are usually more or less uniform in size (but may include rows of smaller scales); 23���32 (n = 15, mean = 26.2, SD = 2.1) medial scale rows on tail in longitudinal series from fracture plane (1 st autotomy septum) to tip; 13���18 (n = 15, mean = 14.4, SD = 1.3) rows of scales across original tail (large row at maximum width); ventral scales considerably smaller than dorsal scales, flat and not arranged in regular rows. Regrown tail: rounded distally, scales not arranged in clear transverse rows. Pattern (in spirit, Fig. 2, 7 - top row). Variable. Most specimens tan to medium-dark brown with heavy, pale spotting on the flanks. Dorsum with an overlay of fine dark reticulations and pale spotting (usually smaller than those on flanks). The vertebral zone has reduced pigmentation and is marked by a series of pale blotches (~ 5���6 between the pectoral and pelvic girdles) which in some specimens are narrowly broken and slightly misaligned across the vertebral line. Head, as for dorsal ground colour with scattered dark flecks or blotches. Canthal stripe absent or weak without sharply defined edges and not contrasting strongly with other facial markings. Limbs with fine reticulations, inner digit of forelimb with reduced pigmentation. Ventral surfaces off-white, immaculate. Original tail as for trunk but usually with broader, dark markings that form a coarse reticulated pattern. Measurements and scale counts of holotype (in mm). QM J 90778 ♀ (Fig. 2), SVL = 60.2, MBW = 14.1, MBD = 7.6, AG = 28.3, L 1 = 17.3, L 2 = 20.0, HL = 11.0, HD = 5.7, HW = 9.1, S = 5.1, EE = 3.3, TL = 23.3, TW = 14.4, scales contacting posterior edge of rostral = 11, scales contacting posterior edge of mental = 10, lamellae beneath 4 th finger = 10, scales beneath 4 th toe (basal junction to tip) = 12, medial scale rows on tail from fracture plane (1 st autotomy septum) to tip = 26, rows of scales across original tail = 14, supralabials = 16, infralabials = 15. Etymology. Named for Dr Andrew Amey for his contributions to documenting Australia���s herpetofauna and promoting access to the Queensland Museum���s reptile and amphibian collections. Comparisons. Diplodactylus ameyi sp. nov. can be distinguished from Diplodactylus barraganae Couper, Oliver & Pepper (in Oliver et. al., 2014), Diplodactylus bilybara Couper, Pepper & Oliver (in Oliver et. al., 2014), D. conspicillatus Lucas & Frost (1897), Diplodactylus custos Couper, Oliver & Pepper (in Oliver et. al., 2014), Diplodactylus hillii Longman (1915) and Diplodactylus laevis Sternfield (1924) in possessing a small first supralabial that is not differentiated from the rest of the supralabial row (vs. enlarged and contacting ventral edge of nasal scale) and in lacking a well-defined canthal stripe (vs. canthal stripe well-defined). It further differs from D. bilybara, D. custos and D. laevis in lacking an acute attenuated extension at the tip of the original tail. Diplodactylus ameyi sp. nov. differs from all known populations of D. platyurus (which may be composite, containing at least two additional candidate taxa - lineages G & H in Oliver et al., 2014) in having a broad Ushaped snout (dorsal view) which is convex along its dorsal edge and lacks a well-defined canthus rostralis (Figs 2, 4 A, 5 A, 6, 7 top row). All` D. platyurus��� populations have finely tapered (V-shaped) snouts with a prominent canthus rostralis, suggesting significant differences in the underlying skull morphology (Figs 1, 4 B, 5 B & C, 7 middle and bottom rows). Diplodactylus ameyi sp. nov. tends to have more scales contacting the posterior edge of the rostral shield (8���13, 10.5 �� 1.3 vs. 5���10, 7.9 �� 1.4 in` D. platyurus���). Diplodactylus ameyi sp. nov. also obtains a greater SVL (max SVL = ~ 60 mm vs. D. platyurus���) (Figure 7). Distribution and vegetation associations (Figs. 8, 9). Diplodactylus ameyi sp. nov. occurs widely across the eastern edge of the Eromanga Basin, from Winton in the north to the north-western edge of New South Wales. The distribution of museum specimens suggests a geographic hiatus between northern and southern samples (Fig. 8) but further fieldwork is required to establish if this is simply an artefact of sampling. This species occurs widely through arid woodland habitats on a range of sand and clay-based substrates throughout its range (typical habitat at one collection site is illustrated in Fig. 9). The following habitat notes are associated with specimens in the Queensland Museum collection: QM J 83467 ���Red sandy soils Acacia cambagei associations���; QM J 59978 ���Open flat, few low bushes. Red sandy soils��� and QM J 74874, QM J 89191 and QM J 90170 were found in association with Mulga woodlands (Acacia aneura). Dietary note. The stomach of QM J 90778 (the holotype, collected 1 May 2010) was examined to determine prey species. Its contents comprised ~ 85 % worker termites and 15 % ants belonging to the genus Iridomyrmex (Formicidae: Dolichoderinae)., Published as part of Couper, Patrick J. & Oliver, Paul M., 2016, A new species of gecko from arid inland regions of eastern Australia (Diplodactylus; Diplodactylidae), pp. 525-538 in Zootaxa 4093 (4) on pages 526-535, DOI: 10.11646/zootaxa.4093.4.4, http://zenodo.org/record/270976

Crenadactylus ocellatus (Gray, 1845) South-western clawless geckos Figs. 4 A, 5, 6, 7 Diplodactylus ocellatus Gray, 1845 Synonymy: Diplodactylus bilineatus Gray, 1845 (fide G��nther 1867) [���South-west��� of Oliver et al. (2010)] Holotype. BMNH 1947.3.6.68 from Western Australia. Specimen obtained from the Buchanan collection from WA, collection date not specified. Synonomy. Diplodactylus bilineatus. Syntypes (2), Gray 1845, BMNH 1947.3.6.69 (2 specimens) from Houtman���s Abrolhos, Western Australia. Collected by ���Mr. Gilbert���, collection date not specified. Diagnosis. A moderately large (to 35.5 mm SVL) species of Crenadactylus with wide head (HW/HL 0.52��� 0.70) and short trunk length (ILL / SVL 0.37���0.50). Rostral in full contact with nostril, internasal (if present) not extending beyond supranasal, 1 or 2 granular postmentals, dorsal scales homogeneous with smooth to weak keels, usually no pre-cloacal pores visible but some males with 2, when present pore-bearing scales in contact at midline, no enlarged tubercles on original tails. Ground colour tan and dark brown; dorsal pattern comprised of poorlydefined longitudinal pale and dark stripes, lateral zones pale with irregular dark stippling, pattern heavily overlain with intermixed pale and dark scales giving an irregular appearance, pale spots or ocelli comprised of 3���6 pale scales usually present in dorsolateral zone. Description. A moderately built, small-bodied gecko (to 35.5 mm SVL); body elongate, oblong in crosssection; head relatively wide (HeadW/HeadL 0.52���0.71), moderately long (HeadL/ SVL 0.27���0.33), moderately depressed (HeadD/HeadL 0.27���0.35); in dorsal view, widest at ear openings; head not distinct from neck with neck only weakly constricted; loreal region convex; snout moderately short (SnL/HeadL 0.32���0.40), tapering to rounded tip; head depth ~1.5���2 times eye diameter; eye moderately large; rostral ~2.5 times wider than high, in broad contact with nostril, dorsal edge concave to accommodate supranasals, rostral groove extends from dorsal edge of rostral to ~50% of rostral height; 0 or 1 small internasals; supranasals horizontally elongate, 2���3 times wider than long; nostrils small, similar in size to postnasals, directed laterally, in contact with rostral, supranasal, 2���4 postnasals and first supralabial; supralabials 7���9, antero-dorso edge of first supralabial in contact with nostril; second supralabial rectangular, wider than high, length equal to first supralabial; supralabials 3���5 slightly smaller than first and second, gradually decreasing in size, all wider than high; scales on crown small and slightly rounded, ~one-third the size of dorsal scales, increasing in size anteriorly onto snout and supraocular regions; scales on snout flat and slightly enlarged, 1.5���2 times larger than scales on crown; supraocular scales enlarged; 1 or 2 soft spines on posterior eyelid; mental triangular to trapezoidal in shape, widest anteriorly, not extending past ventral edge of first infralabial; postmental not enlarged relative to other scales adjacent to infralabials; gulars small near infralabials, decreasing to granular, ~0.1���0.2 the size of scales on ventrum, juxtaposed and slightly rounded; gulars flattened from mental to eye, becoming more rounded from eye to neck; infralabials 7���9, first largest and squareshaped, gradually decreasing in size posteriorly. Limbs short, distance between limbs close to half of SVL (ILL / SVL 0.37���0.50); scales on anterior surfaces of forelimbs elongate, broad and rounded, slightly imbricate, decreasing in size distally, ~2 times the size of scales elsewhere on forelimb; scales on dorsal surface of hind limbs rounded and slightly raised, decreasing in size and becoming more rounded distally; scales on anterior surface of thigh flat and circular, slightly larger in area than scales on hindlimbs elsewhere; fingers 5; toes 5; claws absent from all digits, digits short; fourth finger lamellae in 6 rows; fourth toe lamellae in 8 rows; apical plates much wider than digit; two pairs of dorsal apical plates; one pair of ventral apical plates, round to ovate. Scales on dorsum and lateral surfaces small and homogeneous, smooth to weakly keeled, ~2.5 times wider than deep, juxtaposed; scales slightly raised, highest point at posterior edge of scale, widest at midpoint of scale; scales on ventrum homogeneous, flat, imbricate, ~equal to 1.5 times the size of dorsal scales in area, widest at midpoint, with rounded posterior edge. Tail moderately long, maximum width one-quarter to one-third along tail, gradually tapering to a fine point, round in cross-section; scales on dorsal surface of tail ~2���3 times size compared to dorsum, homogeneous, flattened, imbricate, square to oblong in shape, arranged in transverse rows; ventral tail scales flattened, ~2���3 times the size of dorsal tail scales; single enlarged cloacal spur per side, ~3���4 times the size of surrounding scales; no visible pre-cloacal pores (except on 2 males with 2 pores). Colouration and pattern. Dorsal background colouration tan and dark brown; light brown dorsolateral lines edged with dark brown emanate from back of eye to tail, continues forwards past eye as pale canthal stripe, terminating above nostril; poorly-defined silver-grey vertebral stripe discernible in some specimens, 2���3 scales wide, bordered by poorly defined dark brown paravertebral lines or dark edges of dorsolateral lines; dorsolateral zone with 7���12 widely spaced ocelli between eye and tail, variably defined ocelli cluster comprised of 3���6 pale scales, often lacking or loosely bordered by darker scales; lateral zone with poorly-defined lateral line that extends from postero-ventral edge of eye above limbs to tail, continuing forward as dark loreal stripes to nostril; below lateral stripes on flanks a mixture of darkly pigmented scales with grey and pale scales; crown and snout a mixture of dark brown, brown and pale grey scales; supralabials pale with dark stippling, infralabials moderately pigmented; ventral edge of orbit dark brown; limbs have same intermixture of dark and pale scales with numerous scattered small pale ocelli, weak on front limbs, more prominent on rear; top of digits have a network of dark lines; ventrum pale with fine to heavy dark brown flecking, not forming conspicuous lines; heavier flecking on rear limbs and cloacal region and tail; tail continues dorsal pattern including ocelli on dorso-lateral stripes, dark vertebral zone and light brown paravertebral zone. Distribution. This species occurs in the south-west of Australia (Fig. 2). It is absent from the mesic southwestern corner, extending only as far south-west as Waroona (WAM R88483) then extending eastwards including records from Kojanup, Tambelup and Fitzgerald River National Park. The western-most extent of the distribution is from an old record in Israelite Bay at Cape Arid National Park (WAM R14176) and a more recent record from Balladonia at the edge of the Nullarbor Plain (WAM R157901). It extends into the wheatbelt and to the arid interzone, with records from near Mount Manning and Bowgada Nature Reserves, and an isolated record from Walganna Rock in the north-east (WAM R135103, genotyped as C. ocellatus and shown in Fig. 3 A of Oliver et al. 2010). Occurs on most of the Houtman Abrolhos Islands (off Geraldton) on which surveys have been conducted. Habitat. Found in open woodland habitats throughout its range including areas dominated by Xanthorrhea grass trees, Eucalyptus (jarrah, mallee, marri, tuart, wandoo) and spinifex grass (Triodia) on a wide variety of soft (sandy and loamy) and hard stony substrates (e.g. laterite). Collectors��� notes record specimens have been found under ground cover such as fallen and rotten logs, woodpiles, leaf litter, granite boulders, limestone slabs, sheets of tin, railway sleepers and rubbish. One specimen (WAM R72276) was found one meter high under the bark of a gimlet tree, and other records mention ���under bark��� indicating some climbing ability. Etymology. The specific name ocellatus refers to the pale scattered blotches or ���ocelli��� on the dorsum. Remarks. Our experience with this species indicates they are reasonably common compared to other Crenadactylus species. Within the genus, C. ocellatus is exceptional for its dorsal patterning, with its intermixture of dark and pale scales and less well-defined longitudinal stripes (Fig. 7). In contrast, most other Crendactylus species have relatively ���clean��� longitudinal lines with possibly the exception of C. tuberculatus sp. nov. (Fig. 4 D). It is likely this broken colour pattern may be beneficial for camouflage owing to this species��� ground-dwelling habits. In contrast, the striped patterns in other Crenadactylus taxa are likely beneficial to species that climb in spinifex clumps and are likely convergent with the phasmid geckos in the genus Strophurus Fitzinger which also occur in spinifex., Published as part of Doughty, Paul, Ellis, Ryan J. & Oliver, Paul M., 2016, Many things come in small packages: Revision of the clawless geckos (Crenadactylus: Diplodactylidae) of Australia, pp. 239-278 in Zootaxa 4168 (2) on pages 247-250, DOI: 10.11646/zootaxa.4168.2.2, http://zenodo.org/record/262140, {"references":["Gray, J. E. (1845) Catalogue of the Specimens of Lizards in the Collection of the British Museum. Trustees if the British Museum / Edward Newman, London, xxviii + 289 pp.","Gunther, A. (1867) Additions to the knowledge of Australian reptiles and fishes. Annals and Magazine of Natural History, 20, 45 - 68.","Oliver, P., Adams, M. & Doughty, P. (2010) Molecular evidence for ten species and Oligo-Miocene vicariance within a nominal Australian gecko species (Crenadactylus ocellatus, Diplodactylidae). BMC Evolutionary Biology, 10, 386. http: // dx. doi. org / 10.1186 / 1471 - 2148 - 10 - 386"]}

Oliver, Paul M., Doughty, Paul (2016): Systematic revision of the marbled velvet geckos (Oedura marmorata species complex, Diplodactylidae) from the Australian arid and semi-arid zones. Zootaxa 4088 (2): 151-176, DOI: 10.11646/zootaxa.4088.2.1

Crenadactylus tuberculatus sp. nov. Cape Range clawless geckos Figs. 3 B, 4D, 12, 13 [���Cape Range��� of Oliver et al. (2010)] Holotype. WAM R174304, adult male, Charles Knife Road, Cape Range National Park, North West Cape, Western Australia (22��05'26.48"S, 114��00'39.57"E) by P. Doughty, A.M. Bauer and R.J. Ellis, 19 August 2014. Fixed in 10% formalin, stored in 70% ethanol at WAM. Liver sample stored in 100% ethanol at WAM. Paratypes (6). WAM R61203, Yardie Creek, WA (22��20'S, 113��49'E); WAM R134295, WAM R134627, WAM R134945, 1.5 km SW Vlamingh Head lighthouse, North West Cape, WA (21��48'30"S, 114��06'07"E); WAM R132481, WAM R132483, Shothole Canyon, Cape Range National Park, WA (22��03'S, 114��01'E). Diagnosis. A very small (to 27.7 mm SVL) species of Crenadactylus with long (HL/ SVL 0.30���0.35) and narrow (HW/HL 0.43���0.50) head. Rostral in full contact with nostril, internasal (if present) not extending beyond supranasal, 2 small postmentals, dorsal scales heterogeneous with scattered enlarged keeled tubercles, males with 4 pre-cloacal pores with innermost pore-bearing scales separated by a small intervening scale, no enlarged tubercles on original tails. Ground colour light to dark greyish-brown, dorsal pattern consists of weakly-contrasting pale and dark longitudinal stripes, small pale spots may be present in dorsolateral zone, lateral zones grey with at most faint uniform stippling (not forming irregular lines), ventrum pale off-white with stippling absent to weak. Description of holotype. WAM R174304, an adult male with the following meristics (in mm): SVL 24.8; TailL 11.3 (original); HeadL 7.9; HeadW 3.9; HeadD 2.3; SnL 2.2; ILL 11.2; SupLab 8 (left), 8 (right); InfLab 9, 9; internasals 1; PCP 4. Slender, small-bodied with elongate body, round to oblong in cross section, ~1.5 times wider than deep; head narrow (HeadW/HeadL 0.49), elongate (HeadL/ SVL 0.32), moderately depressed (HeadD/HeadL 0.29); in dorsal view, widest just posterior to eye, slightly wider than widest portion of body; head not distinct from neck, neck weakly constricted; loreal region weakly convex, weakly concave around nasal region; snout elongate (SnL 2.2; SnL/HeadL 0.40), round tip; eye moderately large; rostral ~3 times wider than high, in broad contact with nostril, dorsal edge concave to accommodate first supranasal, rostral groove ~60% of rostral length from dorsal edge below internasal notch; internasal tiny, ~one-fifth the size of first supranasal in area; supranasals two per side; first supranasal (in contact with rostral) oblong, transversely oriented, ~2.5 times wider than high, in broad contact with nostril; second (upper) supranasal small, rounded, ~one-third the size of first supranasal; nostril small, ~one-third size of first supranasal, directed dorsolaterally, in contact with rostral, first and second supranasals, three postnasals, and first supralabial; postnasals 3; supralabials 8 on both sides; antero-dorso edge of first supralabial in contact with nostril, postero-dorso edge notched by third postnasal and first loreal; second supralabial rectangular, wider than high, length equal to length of first supralabial; supralabials 3���8 smaller than first and second, gradually decreasing in size, all wider than high; scales on crown small and flattened, ~equal to size of smaller dorsal scales, ~one-sixth the size of enlarged dorsal tubercles, increasing in size anteriorly onto snout; scales on snout flat, enlarged, ~1.5���2.0 times the size of scales on crown; supraocular scales enlarged; two enlarged conical scales at postero-dorso edge of eye; mental trapezoid-shaped, widest anteriorly, narrowing slightly to posterior edge, terminates just beyond ventral edge of first infralabials; postmentals two, round in shape, ~equal to size of scales posterior to postmental; gulars granular, rounded; infralabials 9 on each side, first largest, square-shaped, gradually decreasing in height and width posteriorly, 1���3 square, 4���9 rectangular, wider than tall. Limbs short, distance between limbs nearly half of SVL (ILL / SVL 0.45), nearly equal to tail length (ILL /TailL 0.99); scales on dorsal surface of forelimbs flattened and rounded, decreasing in size distally; scales on ventral surface of forelimbs similar to dorsal; scales on dorsal surface of hind limbs rounded and slightly raised, decreasing in size and becoming flattened and more rounded distally; scales on ventral surface of hind limbs flat and circular, imbricate, increasing in size distally; fingers 5; toes 5; claws absent from all digits; digits moderately long; fourth finger lamellae in 6 rows; fourth toe lamellae in 8 rows; in dorsal view, enlarged scale between apical plates ~2 times the size of adjacent lateral scales; apical plates much wider than digit; one pair of ventral apical plates, circular, as wide as long. Scales on dorsum heterogeneous: background of small homogeneous scales with scattered enlarged scales with prominent keeling; small dorsal scales homogeneous, slightly raised; enlarged tubercles scattered on dorsum and nuchal region, loosely aligned longitudinally, ~4���6 times the size of surrounding smaller scales in diameter, separated from one another by 1���4 smaller scales, enlarged tubercles have raised midline forming a moderate keel, tallest at posterior edge of scale; some enlarged tubercles in dorso-lateral zone, absent on lateral surfaces; scales on ventrum homogeneous, rounded and flattened, ~2 times the size of smaller scales present on dorsum, all similar in size and shape. Tail original, nearly half of SVL (TailL/ SVL 0.45), tapers gradually from widest point posterior to vent to narrow rounded point; round in cross-section; dorsal scales somewhat rounded and imbricate, variable in size and shape, ~0.25���0.75 the size of enlarged tubercles on dorsum; ventral tail scales similar to those of dorsal tail scales but more flattened; single enlarged post-cloacal spur on both sides of cloaca, ~6 times the size of surrounding scales; pre-cloacal pores 4, forming a partial chevron, pore-bearing scales notched on posterior edge to accommodate pore, two pores on each side separated by a single intervening poreless scale. Colouration and pattern. In life, ground colour light brownish-grey with moderately-contrasting dark brown longitudinal stripes, all stripes with some intermingling of dark and pale scales, some hints of orange on enlarged tubercles. Dark paravertebral stripes enclose a narrow pale vertebral stripe from the crown to posterior to hindlimbs where the paravertebral stripes join; pale dorsolateral stripe from posterior edge of eye to tail, small widely-spaced clusters of orange-tinted enlarged tubercles above hindlimbs and before tail, stripe continuing forwards anterior to eye as pale canthal stripe; above pale canthal stripe a dark triangular patch on snout; crown with a mixture of dark brown and black scales; dark brown dorsolateral stripe from posterior edge of eye to tail, continuing forwards on snout as dark loreal stripe; labial scales pale with dark mottling; lateral surfaces light brown with dark variegations suggesting a lateral line; limbs a mixture of light and dark brown scales with peppering of black scales occasionally forming short lines; ventrum pale off-white with dark stippling on underside of limbs, lower abdomen and between arms, elsewhere lightly stippled; original (see below) tail light to medium brown with scattered black scales. In preservative, colours faded but all stripes discernible. Variation. Most specimens examined conformed to the holotype, with the following exceptions. Enlarged dorsal tubercles tended to not occur anterior to the ear openings, but in some individuals they occurred up to the level of the eyes, especially behind the eyes. Most individuals lacked tails (original or regrown), and we believe the holotype tail is an original based on the scalation and darker colouration compared to regenerated tails in our limited sample. Regrown tails tended to be paler with smaller scales compared to the holotype tail. WAM R134627, a male, has an atypical pattern of three pre-cloacal pores in a single row, but the scales in this area were irregularly-shaped. Females differed from males in that they lacked pre-cloacal pores and the cloacal spurs tended to not protrude laterally as much as those of males. Colouration and pattern. Of the few individuals photographed in life, most of the variation is in shade���from light tan or brown to dark greyish brown. Other variable features are the small clusters of orange-tinted scales within the pale dorsolateral stripe near the legs and base of tail, ranging from absent to conspicuous. Occasionally the upper lateral zone has a dark edge that forms a weakly-defined additional thin lateral stripe, continuing anteriorly from forelimb to ear opening. In contrast to the holotype, the pale spots on the limbs are more conspicuous in most other individuals. Distribution. Restricted to the Cape Range of the North West Cape on the Western Australian coast near Exmouth (Fig. 2). The southernmost records are from Yardie Creek, where it is sympatric with C. occidentalis sp. nov. (Fig. 2). Several records are from Charles Knife Road and Shothole Canyon, with additional records from near Vlamingh Head and the lighthouse at the northern end of the Cape Range. All save one record are from within the boundaries of the Cape Range National Park. Habitat. Most habitat notes indicate collection from living or dead spinifex clumps, and usually in limestone gorges but with some records from coastal dunes. One specimen was collected from a dead log (WAM R88600). The holotype was collected from the upper edge of a rocky gorge with scattered shrubs and spinifex. Etymology. The species name tuberculatus (Latin) is in reference to the enlarged tubercles on the dorsum of this species, unique within Crenadactylus. Remarks. No other Crenadactylus has heterogeneous dorsal scales as observed in C. tuberculatus sp. nov. (Fig. 3 B). The function of these scales is unknown. This species is also Australia���s smallest gecko species (maximum SVL, Published as part of Doughty, Paul, Ellis, Ryan J. & Oliver, Paul M., 2016, Many things come in small packages: Revision of the clawless geckos (Crenadactylus: Diplodactylidae) of Australia, pp. 239-278 in Zootaxa 4168 (2) on pages 256-259, DOI: 10.11646/zootaxa.4168.2.2, http://zenodo.org/record/262140, {"references":["Oliver, P., Adams, M. & Doughty, P. (2010) Molecular evidence for ten species and Oligo-Miocene vicariance within a nominal Australian gecko species (Crenadactylus ocellatus, Diplodactylidae). BMC Evolutionary Biology, 10, 386. http: // dx. doi. org / 10.1186 / 1471 - 2148 - 10 - 386"]}

Oliver, Paul M., Bourke, Gayleen, Pratt, Renae C., Doughty, Paul, Moritz, Craig (2016): Systematics of small Gehyra (Squamata: Gekkonidae) of the southern Kimberley, Western Australia: redescription of G. kimberleyi Börner & Schüttler, 1983 and description of a new restricted range species. Zootaxa 4107 (1): 49-64, DOI: http://doi.org/10.11646/zootaxa.4107.1.2

Crenadactylus pilbarensis sp. nov. Pilbara clawless geckos Figs. 3 F, 4E, 14, 15 [���Pilbara��� of Oliver et al. (2010)] Holotype. WAM R132630, adult female, Burrup Peninsula, Western Australia (20��35'34"S, 116��47'58"E), collected by B. Maryan, 31 May 1998. Fixed in 10% formalin, stored in 70% ethanol at WAM. Liver sample stored at the Evolutionary Biology Unit, South Australian Museum, Adelaide (SAMA). Paratypes (6). WAM R 100988, 80 km south of Telfer, WA (22��20'S, 122��05'E); WAM R132540, Burrup Peninsula, WA (20��36'45"S, 116��47'37"E); WAM R132627, WAM R132629, Burrup Peninsula, WA (20��35'34"S, 116��47'58"E); WAM R 140394, Deepdale Outstation, Robe River Station, WA (21��43'S, 116��11'E) WAM R160059, 55 km east-south-east of Meentheena Outcamp, WA (21��19'19"S, 121��00'08"E). Diagnosis. A small (to 28.4 mm SVL) species of Crenadactylus. Rostral in full contact with nostril, internasal (if present) not extending beyond supranasal, single greatly enlarged postmental, dorsal scales homogeneous and moderately keeled, 6 pre-cloacal pores in males with innermost pore-bearing scales in contact, no enlarged tubercles on original tails. Ground colour light brown; dorsal pattern consists of well-defined pale and dark longitudinal stripes, scattered clusters of 1���3 pale scales, lateral zones light grey to tan with stippling occasionally forming 1 or 2 broken or weakly defined lateral lines; ventrum pale off-white with weak to moderate stippling. Description of holotype. WAM R132630, adult female with the following meristics (mm): SVL 27.0 mm; TailL 24.3 mm (partially regenerated); HeadL 8.4; HeadW 4.1; HeadD 2.5; SnL 3.3; ILL 13.0; SupLab 8 (left), 9 (right); InfLab 8, 9; internasals 1; PCP 0. A slender, small-bodied gecko; body elongate, oblong in cross section, ~1.5���2.0 times wider than deep; head narrow (HeadW/HeadL 0.49), elongate (HeadL/ SVL 0.31), moderately depressed (HeadD/HeadL 0.30); in dorsal view, widest at ear openings, not wider than widest portion of body; head not distinct from neck, neck very weakly constricted; loreal region weakly convex, concave around nasal region; snout elongate (SnL 3.3; SnL/HeadL 0.39), rounded tip; eye moderately large; rostral ~3 times wider than high, in broad contact with nostril, slightly scalloped dorsally to accommodate first supranasal, notched by internasal, rostral groove originates from dorsal edge of scale, ~40% of rostral height; internasal tiny, ~one-fifth the size of first supranasal in area; supranasals 2 per side; first supranasal oblong, transversely oriented, ~2 times wider than high, in broad contact with nostril; second (dorsal) supranasal small, rounded, tapering to blunt point at nostril contact, ~half the size of first supranasal; nostrils small, ~one-third size of first supranasal, directed dorsolaterally, in contact with rostral, first and second supranasals, postnasals, and first supralabial; postnasals 2, equal to size of second supranasal; supralabials 8 on both sides, antero-dorso edge of first supralabial in contact with nostril; second supralabial rectangular, wider than high, length equal to length of first supralabial, supralabials 3���8 smaller than first and second, gradually decreasing in size, all wider than high; scales on crown small and flattened, ~one-third the size of dorsal scales, increasing in size anteriorly onto snout; scales on snout flat, enlarged, ~2���3 times the size of scales on crown; supraocular scales enlarged; single enlarged conical scale at postero-dorso edge of eye; mental trapezoid-shaped, widest anteriorly, narrowing slightly to concave posterior edge, terminates equal to ventral edge of first infralabials; single enlarged postmental, hexagonal in shape, ~2 times area of scales posterior and lateral to enlarged postmental, ~4 times larger than other gular scales; gulars granular, rounded, ~0.25���0.50 the size of scales on ventrum; infralabials 9 on each side, first largest, square-shaped, gradually decreasing in height and width posteriorly, infralabials 1���3 square or rectangular, 4���9 rectangular, wider than high. Limbs short, distance between limbs nearly half of SVL (ILL / SVL 0.48), slightly over half the length of tail (ILL /TailL 0.54); scales on dorsal surface of forelimbs imbricate, flattened and rounded, decreasing in size distally; scales on ventral surface of forelimbs similar to dorsal; scales on dorsal surface of hind limbs rounded and slightly raised, decreasing in size and becoming more rounded distally; scales on ventral surface of hindlimbs flat and circular, slightly larger in area than dorsal hindlimb scales; fingers 5; toes 5; claws absent from all digits; digits moderately long; fourth finger lamellae in 6 rows; fourth toe lamellae in 8 rows; in dorsal view, enlarged scale between apical plates 1.5 times the size of adjacent lateral scales; apical plates much wider than digit; two pairs of dorsal apical plates; one pair of ventral apical plates, circular. Scales on dorsum small and homogeneous, round to oblong, juxtaposed; scales slightly raised, moderately keeled, highest point at posterior edge of scale, widest at midpoint of scale, becoming flatter and imbricate laterally; scales on ventrum homogeneous, flat, imbricate, equal to 1.5 times the size of dorsal scales in area, widest at midpoint, tapering to a rounded or flattened posterior edge. Tail partially regenerated, last 40% of tail regenerated, slightly shorter than SVL (TailL/ SVL 0.90), tapers gradually from widest point posterior to vent, more abruptly in posterior quarter of tail; round in cross-section; scales homogeneous, imbricate and flat, square to oblong in shape; subcaudal scales flattened, ~1.5���2 times the size of dorsal tail scales; single enlarged spur on both sides of cloaca, ~3���4 times the size of surrounding scales. Colouration and pattern. In preservative, pattern comprised of alternating light and dark longitudinal stripes on dorsum and sides; light brown-tan vertebral stripe narrow, 2���4 scales wide, scales lightly stippled with darker brown, originating on back of head, extending length of body, terminates at base of tail where paravertebral stripes merge; paravertebral stripes wider than vertebral, 3���5 scales wide, medium brown, originating from postero-dorsal edge of eye, curving around posterior of head to vertebral stripe, continuing parallel with vertebral stripe for length of body, tapering from hindlimbs to merge and form single broken line on tail; light brown-tan dorso-lateral stripe 3���4 scales wide, extends from postero-dorsal edge of eye, above forelimbs, parallel with vertebral stripe for length of body, over hindlimbs and onto tail, faded and broken on tail; dorsolateral stripe narrow, 3���4 scales wide, ~equal width to paravertebral stripe, dark brown, same as paravertebral, extends from posterior edge of eye above ear opening, above forelimbs along length of body, above hindlimbs and onto tail, ~one-half of tail length; pale lateral zone below dorsolateral stripe with narrow, solid, dark brown lateral stripe between limbs, 2���3 scales wide; weak stripe below lateral stripe on ventro-lateral edge, extending anteriorly from forelimb to ventral edge of ear opening; ventrum pale, heavily stippled with brown flecks; dark triangular patch on snout, crown a mixture of light and dark scales, dark-edged by extension of paravertebral stripes to eyes, top of eyes purplish, dark brown scales below eye on orbit; labials pale, heavily stippled with brown flecks; pale canthal stripe, extension of pale dorsolateral stripe from eye to rostral scale, below a dark loreal stripe, extension of dark dorsolateral stripe; dorsal surface of limbs with mixed pale, light brown and dark brown scales, scattered clusters of pale scales resulting in some spotting, extending onto digits; dorsal stripes (with exception of vertebral) continue to tail but broken up. Variation. Most specimens conformed to the holotype, with some of the following exceptions. Males possessed 6 pre-cloacal pores (one individual with 4) in a continuous row. Pores formed by a notch in the posterior edge of scale. Females lacked pre-cloacal pores. In males the cloacal spurs are larger than those of females and protrude further than for females. Regrown tails tended to be paler with smaller scales compared to the tail of the holotype. The diagnostic character of a greatly enlarged postmental was unambiguous in all specimens examined, although the shape tended to vary from round to oblong (Fig. 3 F). Keeling on the dorsal scales also varied from weak keels to moderately strong. Colouration and pattern. In life, light to dark brown ground colour with contrasting dark and pale longitudinal stripes. In preservative, thin vertebral stripe pale tan-grey, paravertebral stripes to either side emanating from top of eye and joining just before base of tail; pale dorsolateral stripe from antero-dorso portion of eye to tail, continuing anteriorly from eye as pale canthal stripe; dark brown triangular patch on snout continuous with dark brown crown, intermixed with paler scales; dark dorsolateral stripe from ventro-dorso portion of eye to above limbs to tail, anterior to eye forming dark loreal stripes; labials pale, heavily stippled with dark brown; lateral surfaces pale with thin dark brown stripes, uppermost more defined, lower lateral stripe can be extensively broken and indistinct; stippling on pale ventrum variable; limbs a mixture of dark and light brown scales with small clusters of pale scales apparent; regenerated tails a mixture of dark and pale brown scales, forming short lines or showing no pattern. Distribution. Restricted to the Pilbara region of Western Australia (Fig. 2). Most records are from the Pilbara craton, although many specimens have also been collected from the Burrup Peninsula. Absent from offshore islands except Dolphin Island that is separated from the Burrup Peninsula by a channel only several hundred metres wide. Northern records are from Shay Gap and Meentheena Conservation Reserve. The southern-most records are from the Barlee Range in the south-west and Kumarina in the south-east. Outlying records to the east are from Karlamilyi (Rudall River) National Park and Jiggalong. Habitat. Collection records indicate this species occurs in spinifex clumps on rocky hills and in gullies on the mainland, but sandy substrates for the Dolphin Island specimens. Etymology. Named for the Pilbara region where this species occurs., Published as part of Doughty, Paul, Ellis, Ryan J. & Oliver, Paul M., 2016, Many things come in small packages: Revision of the clawless geckos (Crenadactylus: Diplodactylidae) of Australia, pp. 239-278 in Zootaxa 4168 (2) on pages 259-261, DOI: 10.11646/zootaxa.4168.2.2, http://zenodo.org/record/262140, {"references":["Oliver, P., Adams, M. & Doughty, P. (2010) Molecular evidence for ten species and Oligo-Miocene vicariance within a nominal Australian gecko species (Crenadactylus ocellatus, Diplodactylidae). BMC Evolutionary Biology, 10, 386. http: // dx. doi. org / 10.1186 / 1471 - 2148 - 10 - 386"]}

Gehyra girloorloo sp. nov. Kimberley Karst gecko Figs. 4���7 Holotype. WAM R 175045 (field number CCM 3257) (male), Gogo Station, Pillara Range, south entrance of Menyous Gap (18.40439 ��S; 125.83698 ��E), collected on 4 November 2014 by P.M. Oliver, G. Armstrong and P. Skipwith. Paratypes. WAM R 113727 (female), Cadjbut Mine 90 km south-east Fitzroy Crossing (18.7500��S; 126.1500��E); NMV D 77029 and NMV D 77030 (females), and NMV D 77031 and NMV D 77032 (males), Ngumpan Cliff area (18.75625 ��S; 126.06474 ��E); WAM R 175037 (PMO 147) and WAM R 175038 (PMO 149) (males), Diagnosis. Digits broadly expanded basally and subdigital scansors present on all digits of manus and pes. Digit I of manus and pes clawless, penultimate phalanx of digits II���V free from scansorial pad. Differs from non- Australian Gehyra by the combination of: absence of webbing between third and fourth toes, absence of a skin fold along the posterior hindlimb and small adult size (SVL Gehyra by the combination of small body size (SVL Description of holotype. Adult male with following dimensions (in mm): SVL 41.3, HD 4.7, HL 10.2, HW 8.2, HindL 5.4, ArmL 5.0, TrunkL 18.4, NarEye 3.2, SnEye 4.5, InterN 1.5, InterO 3.2, OrbL 2.4, RosW 1.5, RosH 0.9. Summary meristic data are as follows: SupLab 8; InfLab 7; InterNSc 0; PreCloacal 9; FingSubDL 6; ToeSubDL 7. A small (41.3 mm SVL), slender gecko (Fig. 5). Head slightly depressed (HD/HL 0.46), moderately wide (HW/HL 0.80), of similar width to midbody, widest at posterior edge of jaw, tapering to snout, narrows posteriorly to a moderately constricted neck. Snout short (SnEye/HL 0.44), less than twice eye length (OrbL/SnEye 0.54), rounded in dorsal profile, slightly convex from mid-eye to naris when viewed laterally in profile. Eyes moderately sized (OrbL/HL 0.24), pupil vertical. Ear openings small and circular. Nostrils rounded, contacted by rostral, supranasal, two postnasals and first supralabial; second (ventral) postnasal ~ 2 times larger than first (dorsal), supranasals in contact, no internarial scale. Rostral scale rectangular with nearly flat dorsal edge, wider than high, 1.5 mm wide, 0.9 mm high (RosH/RosW 0.58), deeply furrowed, rostral crease extending from dorsal edge ~ 40 % of rostral height. Supralabials 8, infralabials 7. Scales on snout large, up to 3 times larger than scales on crown of head and body, imbricate and slightly projecting along posterior edge; fold of skin across snout from eye to eye (artefact of preservation). A row of slightly enlarged scales above supralabials gradually decrease in size posteriorly. Mental scale triangular, divides postmentals ~ 35 %; postmentals narrowly separated from second infralabial on right side, in point contact on left side; single pair of outer chin shields with curved dorso-postero edge, ~two-thirds the height of postmentals and ~ 30���40 % smaller in area; outer chin shield in point contact with first infralabial on right side, narrowly in contact on left. Body long (TrunkL/SVL 0.46) and slender, slightly depressed and with a weak ventro-lateral fold posterior to axilla. Dorsal midbody scales small and granular, relatively homogenous, conical, juxtaposed; lateral scales slightly larger, more projecting and heterogeneous than those on dorsal surface, slightly imbricate; ventral scales ~ 3 times larger than dorsal scales, flat, diamond-shaped with rounded edges, juxtaposed. Pre-cloacal pores in a series of 9, forming a short continuous chevron with apex orientated anteriorly; pores centred in scales, innermost scales the largest. Two enlarged, rounded and protruding cloacal spurs present on either side of cloaca. Limbs short (ArmL/SVL 0.12; HindL/SVL 0.13); dorsal and ventral scales generally small, rounded and juxtaposed; scales on anterior edge of limbs slightly larger and imbricate. Fingers and toes 5; expanded toe pads oval; distal row of dorsal scales on toe pads elongate and sharply pointed, forming a serrated ���fringe���. Claws long and free, present on digits 2���5, arising from dorsal surface of expanded toe pads and extending above and beyond toe pad, subdigital lamellae divided, 6 pairs on fourth finger, 7 pairs on fourth toe, all pairs of lamellae in contact, apical lamellae undivided and wedge-shaped. Most of tail is original (52.0 mm, TailL/SVL 1.3), very thin, tapering gently to fine tip, slightly dorso-ventrally flattened at base, proximal 31 mm section original, distal 21 mm section regrown. At widest part near base��� 4.3 mm wide, 3.3 mm deep. Dorsal caudal scales small, granular and relatively homogeneous, subcaudal scales in a single large transversely widened series. Colouration of holotype (in preservative). Background colour pale grey, lightly stippled with darker gray, a series of pale circular spots tending to form oblique transverse rows, enclosed by more extensive transverselyoriented darker grey markings or larger variegations; crown of head with small pale spots above tympanum region, snout and sides of head dark grey, no streaks emanating posteriorly from eye, labials heavily stippled; limbs as for dorsum with smaller pale spots; undersurfaces pale off-white; dorsal surface of tail encircled by pale (wider) and dark brown (thinner) bands. Variation. Table 1 presents ranges of variation for the characters measured. In a small number of specimens, the postmental and second infralabial are in point contact, otherwise they are separated. The same pattern holds for internasals���in most specimens the supranasals are in broad contact above the rostral, in a small number a tiny internasal scale is present. Colouration in preservative is as follows: background dorsal colour pale tan or grey (Fig. 6), densely stippled with fine blackish-brown variegations, moderate sized (0.5 ��� 1.5 mm wide) poorly defined pale circular to oval spots scattered on the dorsum, separated by concentrated patches of dark-brown markings that coalesce to form weakly defined transverse bands; density and distribution of dorsal pattern elements varies along body: usually little or no clear pattern on anterior portion of head, more clearly defined on nape with scattered small light spots and indistinct dark blotches, and consisting of alternating series of comparatively large and more well defined pale spots and indistinctly defined transverse brown bands on the torso and original tail. Limbs as for torso, with scattered distinct to indistinct small pale spots; undersurfaces largely unpigmented, but with dense fine dark brown maculations around the ventrolateral regions of head, limbs and ventral tail scales. Regrown tail light greyishbrown with diffuse longitudinally aligned narrow dark streaks. Colouration in life. The following notes are based on photographs of specimens NMV D 77030 and WAM R 175037 shown in Fig. 7 and WAM R 175042 ��� 3 (not shown). Dorsal surfaces pale purplish buff very finely stippled with medium brown, overlain on dorsum and tail with alternating transverse series of pale spots lacking borders and irregular dark-brown blotches that tend to be transversely aligned and intermittently bordered with yellow-orange. Head with smaller pale and dark spots with scattered yellow-orange flecks or fine spots. Regenerated tails often have a distinctive yellowish wash and very fine brown longitudinal markings. Iris dark golden brown with fine black reticulations. Habitat and reproduction. This species is closely associated with dissected limestone outcrops, and also occurs on small trees and shrubs (Fig. 8). It tends to be less common on exposed vertical rock faces than its much larger sympatric congener G. ko i r a Horner. The locally occuring form of Gehyra nana tends to utilise different habitats���it is rarely found on limestone and generally on smaller boulders and rocky debris in surrounding areas. Gehyra girloorloo sp. nov. can be abundant in its preferred habitat, for example, most large shrubs along an open rock face in Menyous Gap had at least one specimen, often positioned head down close to the ground in what appeared to be a hunting posture. In contrast, we found this species to be scarce in early July 2014 when temperatures were low (Gehyra in the variegata-punctata species-group, this species lays a single egg. Some females were gravid in November in spring at the end of the dry season. Distribution. Gehyra girloorloo sp. nov. is only known from a small area of limestone outcropping in the King Leopold Range of the south-west Kimberley on Gogo and Mt Piere Stations (Fig. 1). Known localities extend in a narrow band from the limestone Billy Hills in the north-west, to just north-west of Ngumpan Cliff in the southeast, a straight line distance of approximately 55 km. Etymology. Girloorloo, pronounced ���gir-loor-loo���, is a word the local Gooniyandi mob use for the limestone this species appears to be restricted to. This species name was suggested by elders from the Gooniyandi mob who speak for country on which this species occurs. Comparisons. Gehyra girloorloo sp. nov. can be distinguished from all non-Australian Gehyra by the absence of webbing between third and fourth toes (versus present), the absence of a skin fold along the posterior hindlimb (versus present) and its generally smaller size (max SVL versus > 50 mm). Within Australia, G. girloorloo sp. nov. can be distinguished from all members of the G. australis speciesgroup (which are largely restricted to northern Australia) by its smaller size (max SVL versus > 50 mm), divided subdigital lamellae (versus at least some undivided), lower number of pores in the males (versus > 12) and females laying just one egg per clutch. Gehyra girloorloo sp. nov. can be distinguished from members of the G. variegata group from the Australian arid zone by its lower number of pre-cloacal pores in males (versus usually more than 10 [Hutchinson et al. 2014]), no conspicuous dark streaks emanating from behind the eyes, at most only light red hues (versus rich reddish-brown) and less contrasting dorsal pattern combining diffuse dark markings (versus extensive reticulations) and diffuse light spots (versus tending towards smaller and well defined). A further species in this group, G. pilbara, is similarly small, but differs in dorsal colouration (reddish-brown vs. pinkish-grey), the presence of obviously enlarged loreal scales above the infralabials (versus absent) and in having an extremely short snout resulting in short, wide postmentals (versus tall and thin in G. girloorloo sp. nov.) (Fig 4.). Gehyra girloorloo sp. nov. can be distinguished from most other members of the G. variegata group that also occur in the AMT as follows: from G. xenopus and G. spheniscus by the absence of a wedge of granules between proximal lamellae (versus present), and in the case of the former species, also much smaller size (max SVL 48 versus 79 mm); from G. occidentalis by its lower number of subdigital lamellae (5���7 versus 7���10) and small body size (max SVL 48 mm versus 76 mm); and from G. multiporosa by the absence of dark lateral head streaks, and fewer pores in males (8���11 versus 20���49). A final taxon from this region, G. nana, is a complex of species (unpublished data). However, G. girloorloo sp. nov. can be distinguished from both geographically proximate and type G. nana by its pinkish-grey dorsal colouration (versus reddish), background stippled (versus plain), larger and more diffuse pale spots (versus small and clearly defined), diffuse and transversely-oriented dark brown blotches on dorsum (versus clearly defined brown blotches or [usually] spots), and low number of pre-cloacal pores in males (8���11 versus 11���17). Gehyra nana from around localities from where G. girloorloo sp. nov. has been recorded are also particularly small (SVL 39.0, 35.0��� 41.9 mm) and strongly spotted (see Fig. 5)., Published as part of Oliver, Paul M., Bourke, Gayleen, Pratt, Renae C., Doughty, Paul & Moritz, Craig, 2016, Systematics of small Gehyra (Squamata: Gekkonidae) of the southern Kimberley, Western Australia: redescription of G. kimberleyi B��rner & Sch��ttler, 1983 and description of a new restricted range species, pp. 49-64 in Zootaxa 4107 (1) on pages 56-61, DOI: 10.11646/zootaxa.4107.1.2, http://zenodo.org/record/260932, {"references":["Hutchinson, M. N., Sistrom, M. J., Donnellan, S. C. & Hutchinson, R. G. (2014) Taxonomic revision of the Australian arid zone lizards Gehyra variegata and G. montium (Squamata, Gekkonidae) with description of three new species. Zootaxa, 3814 (2), 221 - 241. http: // dx. doi. org / 10.11646 / zootaxa. 3814.2.4"]}

Oedura cincta de Vis, 1888 Inland marbled velvet gecko Figs. 3, 5���6 Lectotype. QM J226, Charleville, Queensland, Australia (collected by Kendall Broadbent in 1885). Paralectotypes. AMS R5602, AMS R5603, as for lectotype. Synonomy. Oedura derelicta Wells & Wellington, 1985, holotype���NTM R11413, Jessie Gap, Alice Springs, Northern Territory. Diagnosis. A large-bodied (SVL: mean 90 mm, max 108 mm) species in the O. marmorata complex, with a moderately broad head (HW/SVL 0.17���0.21), moderately long body length (Trk/SVL 0.42���0.53), moderately long tail (original TL/SVL 0.61���0.79) narrower than head and roughly circular in cross-section, rostral usually completely divided, apical lamellae wide (ToeW/SVL 0.023���0.036), subdigital lamellae in a flared series that on fingers 3 and 4 is wider than apical pair, 9���21 precloacal pores in adult males and dorsal pattern (especially conspicuous on juveniles) of 5 or 6 light transverse bands on purplish-brown background. Description. A large (max SVL 108 mm) and moderately elongate Oedura (Trk/SVL 0.42���0.53). Head moderately wide (HW/SVL 0.17���0.21) and deep (HD/SVL 0.07���0.12). Rostral usually fully (n = 39) or near fully divided (n = 5), bordered dorsally by two internasals, internasal bordered dorsally by two relatively small, roundish supranasals and 3���5 (mode = 3) small intervening scales. Enlarged supralabials 9���11 to midpoint of eye, 9���13 in total, second supralabial equal to or slightly higher than first, infralabials 10���15. Limbs moderately long (ArmL/ SVL 0.11���0.14, LegL/SVL 0.12���0.16). Subdigital lamellae expanded and prominent, 6���9 under third finger, 8���10 under third toe, distal lamellae on digits either divided or deeply notched, apical pair distinctly larger (but not always wider) than and separated from other pairs, penultimate pair slightly narrower, subsequent lamellae on digits two to five forming distinctly flared series, proximal lamellae undivided. Enlarged postcloacal spurs in a single cluster of between 1���4 (mode 2). Original tail moderately long (TL/SVL 0.58���0.79) and narrow (TW/SVL 0.07���0.15) with a slight ventral groove, varying from somewhat depressed to almost circular in cross section, width and depth varies greatly depending on body condition, caudal scalation homogeneous, squarish flat scales arranged in regular rows. Fully regrown tails shorter (TL/SVL 0.43���0.65) than original tails but comparably wide (TW/SVL 0.09���0.13). In preservative, base colouration of dorsum usually dark purplish brown, with a nuchal band and 5 further narrow pale buff transverse bands between the limbs, and a further 5���8 similar bands on the tail. Pale bands often with dark brown regions within them. Nuchal band usually joins a light lateral band extending from the supralabial scales and above the tympanum. Bands usually become indistinct with increasing size, especially on the tail, and are not or only faintly apparent on larger specimens, especially the Central lineage. Extensive further light flecks and blotches are also present between the bands, and elsewhere on the dorsal and lateral surfaces of the torso, limbs and tail. Larger specimens can be heavily mottled with equal amounts of light buff and brown, with no obvious transverse banding. Top of head usually with light to heavy pale mottling and a distinct to indistinct canthal stripe. Venter relatively plain light buff with a faint to distinct wash of light brown, particularly towards the lateral extremities and under head and limbs. Regrown tails dark brown and mottled with varying amounts of light flecking. Hatchlings and juveniles lack light flecking and usually have a very simple pattern of sharply defined light bands separated by wide plain dark brown regions, and a thin white labial stripe. As specimens increase in size, the sharp definition of light and dark regions decreases, and the amount of light flecking and blotching across the body increases. In life, adults possess base colouration of dark purplish-brown during the day, washed out grey at night; light bands and flecks tend to be light grey or buff in the Eastern population, but tend to be more yellow in Central population. Iris dark brown. Particulars of syntypes. QM J226, female with original tail (in mm): SVL 88.0, HW 15.3, HD 9.8, HL 19.2, EN 6.6, IN 2.9, IO 6.9, EYE 4.4, TrK 39.3, ArmL 9.9, LegL 11.4, TL 66.0, TW 12.4, TD 9.8, 3FW 2.4, 3TW 2.7. Scales: RC 100%, SuL 9 (10), InF 12, CS 2/3, 3FL 8, 3TL 8. AMS R5602, male with original tail: SVL 79.0, HW 15.8, HD 9.8, HL 18.5, EN 6.4, IN 2.7, IO 6.8, EYE 4.5, Trk 39.4, ArmL 10.5, LegL 12.2, TL 56.0, TW 11.0, TD 8.1, 3FW 2.2, 3TW 2.2. Scales: RC 100%, SuL 10(10), InF 12, CS 2/2, 3FL 7, 3TL 8, precloacal pores 23. AMS R5603, male with original tail: SVL 78, HW 16.0, HD 9.7, HL 18.8, EN 6.8, IN 2.9, IO 7.3, EYE 4.6, TrK 39.1, ArmL 10.2, LegL 11.9, TL 62.0, TW 8.7, TD 9.0, 3FW 2.3, 3TW 2.7. Scales: RC 100%, SuL 11 (12), InF 12, CS 2/2, 3FL 8, 3TL 8, precloacal pores 11. All three syntypes have original tails and completely divided rostrals. Colour pattern is greatly faded, although indistinct dark bands, blotches and flecks are faintly discernable to a varying extent across the dorsal surfaces of the head, torso and especially tail. Distribution and habitat. The Eastern population occurs in woodlands throughout the channels and floodplains of the Cooper and Darling Basins in western Queensland and New South Wales. Apparently isolated populations occur from the northern Flinders Ranges and just north of the Murray River in South Australia (Fig. 2). Eastern animals are usually associated with exfoliating bark and hollows in small trees (both living and dead), especially Bulloaks in the genus Casuarina, but may also use rocks when available (Miller 1980; P.M. Oliver, pers. obs.; D. Armstrong, pers. comm.). In areas with a high density of suitable trees O. cincta can be abundant, and large numbers can be collected in small periods of time. Specimens have also been reported from under exfoliating rock in the Flinders Ranges (Miller 1980; D. Armstrong, pers. comm.). The Central population is known from two apparently isolated subpopulations centred on the MacDonnell and Reynolds Ranges in the southern Northern Territory. The accuracy of an additional old record (Museum Victoria) from the Tennant Creek area requires verification through additional recent samples. Populations of this species from the Central Ranges are associated with gorges and other range country and are usually found while foraging on and around rock faces. Comparisons. Oedura cincta is very similar the Western lineage (see below) and shares with that taxon a combination of moderately large size (average SVL> 90 mm SVL), moderately long tail which tends to be rounded in cross-section and not wider than the head, prominently flared subdigital lamellae series (on fingers 3 and 4 of equal or greater width to the apical lamellae) and dorsal pattern usually consisting of numerous poorlydefined light flecks and blotches, and often thin light bands. Oedura cincta, however, has a longer rostral crease (60���100% versus 25���60% of the rostral height in the Western lineage). Oedura cincta can be specifically distinguished from lineages in the O. marmorata species complex from northern Australia by possessing a longer (TL/SVL 0.65���0.80 versus 0.53���0.63) and narrower tail (always narrower than the head), a longer body (Trk/SVL 0.42���0.53 versus 0.39���0.49), slightly wider apical lamellae (0.23���0.36 versus 0.21���0.31) and a distinctly flared subdigital lamellae series on fingers 3���4 wider than the apical pair (versus not wider). Oedura cincta can be specifically distinguished from most other Oedura from eastern Australia by the presence of a subdigital lamellae series on digits 2���5 that is prominently expanded at its midpoint (versus tapering or slightly expanded), higher number of postcloacal tubercles per side (mode of 2 versus 1) and dorsal colour pattern consisting of 5���6 distinct to indistinct narrow light bands with poorly defined light flecking (versus wide pale transverse bands in O. castelnaui, distinct dark-edged ocelli or transverse bands of varying size in O. coggeri Bustard, 1966, O. monilis and O. tryoni de Vis, 1884, or two pale bands across the nape and base of tail in O. jowalbinna Hoskin & Higgie, 2008). Further distinguished from the Gulf lineage by larger body size (SVL 78���106 mm versus 64���92 mm), longer original tail (TL/SVL 0.58���0.79 versus 0.49���0.65), wider terminal lamellae and wider lamellae series on fingers (see details below)., Published as part of Oliver, Paul M. & Doughty, Paul, 2016, Systematic revision of the marbled velvet geckos (Oedura marmorata species complex, Diplodactylidae) from the Australian arid and semi-arid zones, pp. 151-176 in Zootaxa 4088 (2) on pages 162-165, DOI: 10.11646/zootaxa.4088.2.1, http://zenodo.org/record/258925

Crenadactylus naso Storr, 1978 Crenadactylus naso species complex Northern clawless geckos Figs. 4 G, 4H, 18, 19 Crenadactylus ocellatus naso Storr, 1978 [���Kimberley B���G��� of Oliver et al. (2010, 2012b)] Holotype. WAM R56206, adult male, Crystal Creek, Western Australia (14��30'S, 125��47'E), collected by L.A. Smith and R.E. Johnstone, 2 November 1976. Fixed in 10% formalin, stored in 70% ethanol at WAM. Paratypes (8). WAM R41373 ���74, Heywood Island, Bonaparte Archipelago, WA (15��20'S, 124��20'E); WAM R43220���21, WAM R43224, Mitchell Plateau, WA (14��57'S, 124��20'E); WAM R56185 ���87, Crystal Creek, WA (14��30'S, 125��47'E). Diagnosis. A small (to 30.7 mm SVL), slender Crenadactylus. Rostral usually in narrow contact with nostril, sometimes excluded (especially in eastern Kimberley), internasals not extending beyond supranasals, gular scales granular, dorsal scales homogeneous and smooth to weakly keeled, 4���8 pre-cloacal pores in contact and forming gradual curve, enlarged tubercles on original tails. Ground colour light grey to dark brown; dorsal pattern consists of weak to well-defined pale and dark longitudinal stripes, dark stripes with scattered single pale scales; lateral zone light grey to tan, ventrum pale grey to off-white, lateral zone and ventrum immaculate to moderately stippled with occasional thin longitudinal lines. Description of holotype. WAM R56206, adult male with the following meristics (in mm) and counts: SVL 26.8; TailL 22.2 (detached, partially regenerated); HeadL 7.7; HeadW 4.1; HeadD 2.5; SnL 3.3; ILL 11.6; SupLab10 (left), 9 (right); InfLab 9, 9; internasals 3; PCP 6. A slender, small-bodied gecko; body elongate, oblong in cross section, ~1.5���2.0 times wider than deep; head narrow (HeadW/HeadL 0.53), elongate (HeadL/ SVL 0.28), moderately depressed (HeadD/HeadL 0.32); in dorsal view, widest at ear openings, not wider than widest portion of body; head not distinct from neck, neck weakly constricted; loreal region weakly convex, concave around nasal region; snout elongate (SnL 3.3; SnL/HeadL 0.43), rounded tip; eye moderately large; rostral ~3.5 times wider than high, in narrow contact with nostril, dorsal edge faintly scalloped to accommodate first supranasals, rostral groove absent; internasals 3, variable in size, ~0.5���0.75 the size of first supranasals in area; supranasals two per side; supranasals on left side, first oblong, transverselyoriented, wider than long, ~1.5 times wider than long, in broad contact with nostril, second round, ~half the size of first, in broad contact with nostril; supranasals on right side, first oblong, transversely oriented, wider than long, ~1.75 times wider than long, in broad contact with nostril, second oblong, transversely-orientated, equal in size to first, in broad contact with nostril; nostrils small, directed dorsolaterally, in contact with rostral, first and second supranasals, postnasals and first supralabial; postnasals 3, oblong in shape, longer than high; supralabials 10 (left), 9 (right), antero-dorso edge of first supralabial in contact with nostril; second supralabial rectangular, wider than high, slightly longer than first supralabial, supralabials 3���10 (left)/9 (right) smaller than first and second, gradually decreasing in size, all wider than high; scales on crown small and flattened, ~half the size of dorsal scales, increasing in size anteriorly onto snout; scales on snout flat, enlarged, ~2���3 times the size of scales on crown; supraocular scales enlarged; two enlarged conical scales at postero-dorso edge of eye; mental trapezoid-shaped, widest anteriorly, narrowing slightly to concave posterior edge, posterior width ~one-third of anterior width; gulars granular, rounded, ~0.50���0.75 the size of scales on ventrum; infralabials 9 on each side, first largest, squareshaped, gradually decreasing in width and height posteriorly. Limbs short, distance between limbs nearly half of SVL (ILL / SVL 0.43), slightly over half the length of tail (ILL /TailL 0.52); scales on dorsal surface of forelimbs imbricate, rounded and slightly raised, becoming flatter distally; scales on ventral surface of forelimbs similar to dorsal; scales on dorsal surface of hindlimbs rounded and slightly raised; scales on ventral surface of hind limbs rounded and slightly raised, becoming flatter distally; fingers 5; toes 5; claws absent from all digits; digits moderately long; fourth finger lamellae in 7 rows; fourth toe lamellae 8 rows; in dorsal view, enlarged scale between apical plates ~2 times the size of adjacent lateral scales, apical plates much wider than digit; one pair of ventral apical plates, circular. Scales on dorsum small and homogeneous, round to oblong, widest at midpoint of scale, becoming flatter and imbricate laterally, juxtaposed; dorsal scales slightly raised, highest point at posterior edge of scale, weakly keeled; scales on ventrum homogeneous, flat, imbricate, equal to 1.5 times the size of dorsal scales in area, widest at midpoint, rounded posteriorly. Tail detached, partially regenerated, last 60% of tail regenerated, slightly shorter than SVL (TailL/ SVL 0.82), tapers gradually from widest point posterior to vent; round in cross-section; scales rounded and slightly raised, imbricate, 2 longitudinal rows of enlarged tubercles along dorsolateral edges of tail, similar in area to surrounding scales, but more prominently raised; ventral tail scales flattened and imbricate, ~equal to 1.5 times the size of dorsal tail scales; scales on regenerated portion rounded and slightly raised on dorsal surface, enlarged tubercles absent, ventral tail scales flat and imbricate; single enlarged raised post-cloacal spur on both sides of cloaca, ~1.5��� 2 times the size of surrounding scales; pre-cloacal pores 6, arranged in a continuous gradual curve, innermost porebearing scales in contact, pore-bearing scales notched on posterior edge to accommodate pore. Colouration and pattern. In preservative, colouration faded to medium brown, pattern comprised of alternating pale and dark longitudinal lines; pale vertebral stripe bordered by dark brown; medium brown paravertebral zone; pale dorsolateral stripes bordered with dark brown; pale and brown lateral stripes somewhat discernible; crown of head pale; top of snout dark brown with pale canthal stripe (extension of pale dorsolateral stripe anterior to eye) and dark brown loreal stripe (extension of paravertebral zone); labial scales pale; arms with feint longitudinal lines; hindlimbs with longitudinal dark and pale stripes; ventral surface light brown; original portion of tail with feint dark and pale stripes, ventral surface light brown; regenerated portion uniform light brown. Variation. The holotype specimen is somewhat faded and shrivelled, whereas more recently-collected material has more vivid colouration and is in better condition. Within the complex, body size to 30.7 mm SVL, ILL / SVL 39���54%; the rostral-nostril contact varies from narrow to point contact and also exclude in some individuals; rostral groove varied from 0���40%, internasals usually 2 or 3 (occasionally 1 or 4), SupLab and InfLab 7���10; precloacal pores usually 6 (occasionally 4 or 8), innermost pore-bearing scales in contact. Dorsal scales homogeneous, smooth to weakly keeled. In life, light yellowish-grey background colour with alternating pale and dark longitudinal lines; pale vertebral stripe often forks on back of head; pale dorsolateral zone with small spots of 1���2 pale scales dotted at regular intervals and continuing on to tail on enlarged tubercles. In preservative, pale vertebral stripe, bordered by medium brown paravertebral stripes with a thin dark brown border, pale dorsolateral zone originates on postero-dorsal edge of eye and continues on to tail, below dorso-lateral zone is a brown lateral zone with dark border, greyish-brown weakly defined lateral stripe between limbs, dorsum has variably expressed small isolated scattered pale scales, sometimes bordered by dark brown scales forming small ocelli, limbs dark medium dark brown with scattered groups of pales scales forming spots, snout dark brown, pale canthal and dark loreal stripes prominent; labials heavily stippled; ventrum pale, finely stippled, especially on chest and neck region, medium peppering of dark brown, sometimes forming weak longitudinal lines; pale enlarged tubercles on tail, occasionally with black scales adjacent. Distribution. As defined here, this species complex occurs throughout most of the Kimberley, except for the Devonian Reef ranges in the south-west which are occupied by C. rostralis, and a small number of other records of uncertain status from the southern Kimberley (Fig. 2). We tentatively assign populations from the Northern Territory and Queensland to this taxon as well. Habitat. Collected from rocky areas with spinifex, including screes, sandstone outcrops and gorges. Numerous specimens were collected by burning spinifex, with occasional records from vine thickets and more open woodland. Etymology. The word naso (Latin) means ���long-nosed��� in reference to the long snout (Storr et al. 1983). Remarks. Despite extensive searches for diagnostic characters among lineages in the C. naso species complex, we could not find non-overlapping characters amongst the few existing voucher specimens. Rather than delay the resolution of the southern and arid zone taxa, by redescribing C. rostralis we have hopefully isolated the remaining taxonomic issues within Crenadactylus to a monophyletic unit. The resolution of the C. naso species complex may be possible through examination of internal characters, and would benefit from a larger series of voucher specimens (with photos in life) to better understand patterns of morphological variation in the group., Published as part of Doughty, Paul, Ellis, Ryan J. & Oliver, Paul M., 2016, Many things come in small packages: Revision of the clawless geckos (Crenadactylus: Diplodactylidae) of Australia, pp. 239-278 in Zootaxa 4168 (2) on pages 265-267, DOI: 10.11646/zootaxa.4168.2.2, http://zenodo.org/record/262140, {"references":["Storr, G. M. (1978) Seven new gekkonid lizards from Western Australia. Records of the Western Australian Museum, 8, 337 - 352.","Oliver, P., Adams, M. & Doughty, P. (2010) Molecular evidence for ten species and Oligo-Miocene vicariance within a nominal Australian gecko species (Crenadactylus ocellatus, Diplodactylidae). BMC Evolutionary Biology, 10, 386. http: // dx. doi. org / 10.1186 / 1471 - 2148 - 10 - 386","Oliver, P. M., Doughty, P. & Palmer, R. (2012 b) Hidden biodiversity in rare northern Australian vertebrates: the case of the clawless gecko (Crenadactylus, Diplodactylidae) of the Kimberley. Wildlife Research, 39, 429 - 435. http: // dx. doi. org / 10.1071 / WR 12024"]}

Iannella, Amy, Oliver, Paul, Richards, Stephen (2015): Two new species of Choerophryne (Anura, Microhylidae) from the northern versant of Papua New Guinea's central cordillera. Zootaxa 4058 (3): 332-340, DOI: http://dx.doi.org/10.11646/zootaxa.4058.3.2

Choerophryne grylloides sp. nov. (Fig. 1 B) Holotype. SAMA R 67921 (Field No. SJR 12700), adult male, calling when collected, un-named camp in upper Sepik River basin, West Sepik Province, Papua New Guinea (near 4 �� 39.185 ���S, 141 �� 43.448 ���E; 900 m a.s.l), collected by S. Richards 0 2 December 2009. Referred specimen. A single specimen, SAMA R 67928 (Field No. SJR 12885), un-named camp in upper Sepik River basin, West Sepik Province, Papua New Guinea (4 �� 37.247 ���S, 141 �� 41.380 ���E; 440 m a.s.l), collected by S. Richards on 13 December 2009, is tentatively assigned to this species given its morphological similarity to the holotype and the proximity of the collection site to the type locality. However, as no call or genetic data are available, this specimen is not nominated as a paratype. Diagnosis. A small species of Choerophryne (SUL 12.5 mm) with a distinctly projecting, elongated snout (OHG/SUL 0.084); urostyle projecting posteriorly beyond the insertion of the hind limbs; long legs (TL/SUL 0.42); expanded discs present on all digits except the first finger, but not more than 1.4 times width of penultimate phalanx; dorsum densely tuberculate; and advertisement call consisting of 4���5 (normally five) distinctly pulsed notes, of which the last comprises more than double the number of pulses of its predecessors (9���10 vs. 3���4). Comparison with other species. Choerophryne grylloides sp. nov. differs from all congeners previously included in the genus Albericus by having a distinctly projecting snout comprising an extension of the nasal bones and the alary processes of the premaxillae. From other long-nosed Choerophryne it differs in the following characters: it is much smaller (SUL C. bryonopsis, C. longirostris, C. microps, C. nigrescens and C. proboscidea (SUL ��� 15 mm) and further differs from these species in lacking an expanded digital disk on the first finger. It is slightly smaller than C. epirrhina sp. nov., C. gracilirostris and C. rostellifer (male SUL 12.5 mm vs. 14.9 ���15.0, 13.5���14.7 and 14.0��� 16.8 mm respectively); and further differs from C. gracilirostris and C. rostellifer in having dense dorsal tuberculation (tubercules much smaller and less densely arrayed in C. rostellifer, localised to the flanks in C. gracilirostris). Choerophryne grylloides sp. nov. is distinguishable from most other small long-nosed Choerophryne species (C. allisoni, C. amomani and C. burtoni) by its longer snout (OHG/SUL 0.084 vs. ��� 0.074) and longer tibiae (TL/ SUL 0.42 vs. ��� 0.38), and from C. arndtorum by its slightly longer tibiae (Tl/SUL 0.42 vs. 0.38���0.42) and slightly shorter head (HL/SUL 0.33���0.34 vs. 0.34���0.39). Key diagnostic features of long-nosed Choerophryne found on the central cordillera of New Guinea are summarised in Table 2. In having calls normally containing five notes Choerophryne grylloides sp. nov. differs from C. amomani (1���2 notes), C. bryonopsis (1 note), C. epirrhina sp. nov. (4 notes), C. longirostris (3���4 notes), C. nigrescens (6���10 notes), C. proboscidea (3 notes) and C. rostellifer (3���4 notes). From all long-nosed species that produce 5 -note calls C. grylloides sp. nov. differs in having terminal notes containing many more pulses than the non-terminal notes (9���10 vs. 3���4). Species regularly or rarely producing 5 -note calls further differ in the following characters (comparisons with non-terminal notes of C. grylloides sp. nov. only): Choerophryne arndtorum has much longer notes (0.13���0.18 vs. 0.04��� 0.06 s) produced at a slower rate (2.7���3.4 /s vs. 15.7���17.5 /s); C. gracilirostris produces call notes at a slower rate (7.2���13.3 /s vs. 15.7���17.5 /s) and containing more pulses (4���9 vs. 3���4); C. burtoni notes are much longer (0.18��� 0.29 s vs. 0.017���0.035), contain many more pulses (9���15 vs. 3���4) and are produced at a much slower rate (1.36���1.64 /s vs. 15.7���17.5 /s); and Choerophryne microps produces 5���9 notes calls with much longer notes (0.055���0.105 vs. 0.017��� 0.035 s) produced at a much slower rate (7.1 /s vs. 15.7���17.5 /s). Description of holotype. Adult male with a small ventral incision on the right side, liver removed, and with the following measurements (in mm): SUL 12.5; TL 5.3; HW 4.3; HL 4.2; ED 1.3; END 1.9; IND 0.9; SL 2.9; OHG 1.1; F 1 D 0.1; F 3 D 0.5; T 1 D 0.3; T 4 D 0.6. Head rather narrow (HW/SUL 0.34), canthus rostralis weakly defined, smoothly rounded, straight in dorsal view, loreal region very slightly concave. Snout slender, projecting well beyond anterior margin of lower jaw (OHG 1.1 mm), tip rounded in dorsal and lateral profiles, ventral surface of tip with an unpigmented medial ridge, widest anteriorly and tapering towards anterior margin of the jaw. Nares oval, located close to tip of snout, oriented ventrolaterally, not visible in dorsal view, internarial distance moderately large (IN/SUL 0.073). Eye moderately small (ED/SUL 0.10), transverse diameter less than length of snout, pupil horizontal. Tympanic annulus indistinct, ventral edge faintly apparent. Tongue oval, two slightly curved prepharyngeal ridges present; vomerine teeth absent, vocal slits present just anterolateral to base of tongue. Urostyle projecting beyond vent, tip rounded and slightly elevated. Dorsum, flanks and dorsal surface of hindlimbs with scattered low, rounded tubercules; venter smooth. Fingers moderately long and slender, with distinct circum-marginal grooves, relative lengths III>IV���II>I, finger I more than half length of II, fingers II���IV with distinctly expanded discs up to 1.4 times width of penultimate phalanx, finger I lacking expanded disk and with only faint circum-marginal groove; subarticular and metacarpal tubercles low and poorly defined, webbing absent. Legs moderately long (TL/SUL 0.42). Toes long and slender, relative lengths IV>III>V>II>I, all toes with expanded terminal discs up to 1.4 times width of penultimate phalanx, all with circum-marginal grooves; metatarsal tubercles and webbing absent. Disks on toes wider than those on fingers (F 3 /T 4 0.78). In preservative, ground colour of dorsum dark brown grading laterally to medium brown, overlain with dense darker brown maculations, slightly denser anteriorly than posteriorly. Snout pale tan with few dark-brown maculations. Inguinal regions with a pair of indistinctly edged white spots. Vent bordered dorsally by a pale-buff posterior band and ventrally by a very-dark-brown band. Exposed dorsal and lateral surfaces of limbs medium brown, extensively flecked with varying darker brown and unpigmented blotches; darker pigmentation on limbs tends not to form distinct patterns, with the exception of some indistinct very-dark-brown transverse bands at the midpoint of thigh, shank and foot. Dorsal surfaces of hands and feet medium brown with extensive darker-brown flecking and numerous unpigmented patches that are increasingly large towards the distal extremities. Venter pale buff with fine tan maculations coalescing on the throat, thighs and ventral surface of limbs and feet. Palmar and plantar surfaces largely unpigmented, with light-tan speckling, coalescing to form a darker brown band at the base of the distal phalanges. Variation. Referred specimen SAMA R 67928 is slightly smaller than the holotype, with slightly shorter tibiae (TL/SUL 0.51 vs. 0.42), but is otherwise of similar dimensions (Table 1). In preservative, dorsal colouration of the referred specimen is much paler than the holotype, being predominantly pale grey with a dark-brown, roughly Xshaped marking spanning the dorsum. Limbs and venter have similar colouration to the holotype. Dorsum less densely tuberculate than holotype. Appearance in life. A photograph in life (Figure 1 B) of the holotype taken at night, shows stronger colouration and sharper tonal variation than is evident in preservative. Medial area of dorsum black, strongly contrasting with orangish-brown snout, thighs and posterior of dorsum. Flanks and limbs with extensive scattered pale grey-blue spots. Iris black with extensive small orange-bronze flecks. Pupil horizontal, encircled by a distinctive thin bright-orange ring. Dorsal surfaces of body densely tuberculate. Advertisement call. A single complete call from the holotype is presented in Figure 2 B. A call bout containing nine consecutive calls produced by the holotype (SAMA R 67921) lasted for 9.9 seconds, and calls were produced at a rate of 0.8 calls/s. The dominant frequency was very high, ranging from 4845 hz to 5115 hz (mean = 4978, SD = 104; n = 7 calls). Calls contained 4���5 notes (but eight of nine calls had 5 notes), and were 0.23��� 0.30 s long (mean = 0.28, SD = 0.02), narrowing to 0.29��� 0.30 s (mean = 0.29, SD = 0.005) when the 4 -note call was excluded. Non-terminal notes of each call contained 3���4 pulses ranging in length from 0.017��� 0.035 s (mean = 0.023, SD = 0.035; n = 27). Notes were produced at a rate of 15.7���17.5 notes/s (mean = 16.5, SD = 0.5; n = 9). Pulse rate in non-terminal notes was 93.8���130.4 /s (mean = 113.5, SD = 9.7). Terminal notes in each call were longer (0.043��� 0.056 s, mean = 0.049, SD = 0.004; n = 7 calls) and contained a higher number of pulses (9���10 vs. 3���4) produced at a faster rate (167���212 pulses/s, mean = 185, SD = 14.0; n = 7 notes). Etymology. The specific name is a feminine compound adjective, comprising the Latin gryllus (cricket) and the Greek - oides (resembles), alluding to the resemblance of the high-pitched advertisement calls of this species to the stridulatory sounds made by crickets. Natural history and distribution. C. grylloides sp nov occurs in primary foothill rainforest on the northern side of Papua New Guinea���s central cordillera at altitudes of 440���900 m a.s.l. The holotype was calling at night from the exposed surface of a leaf, roughly 30 cm from the ground. No other individuals were seen or heard at the type locality; however, given the extent of suitable habitat in the area, this species probably has a broad distribution along the northern slopes of the central cordillera., Published as part of Iannella, Amy, Oliver, Paul & Richards, Stephen, 2015, Two new species of Choerophryne (Anura, Microhylidae) from the northern versant of Papua New Guinea's central cordillera, pp. 332-340 in Zootaxa 4058 (3) on pages 337-339, DOI: 10.11646/zootaxa.4058.3.2, http://zenodo.org/record/244825

Choerophryne epirrhina sp. nov. (Figs 1 A, 2) Holotype. SAMA R 67920 (Field No. SJR 12687), adult male, calling when collected, un-named camp in upper Sepik River basin, West Sepik Province, Papua New Guinea (4 �� 39.185 ���S, 141 �� 43.448 ���E; 850 m a.s.l.), collected by S. Richards 01/ 12 / 2009. Paratopotype. Field No. SJR 12684, adult male, with same locality and collector details as holotype. To be deposited in PNGNM. Diagnosis. A moderately small Choerophryne (SUL 14.9 ���15.0 mm) with a projecting and elongated snout (OHG/SUL 0.09���0.10); moderately long legs (TL/SUL 0.41); expanded discs up to 1.7 times width of penultimate phalanx on all digits of hands and feet except the first; and advertisement call consisting of 3���4 (normally 4) distinctly pulsed notes repeated at a rate of 2.2���2.3 /s and comprising 27���31 pulses produced at a rate of 117���141 /s. Comparison with other species. Choerophryne epirrhina sp. nov. differs from all congeners previously included in the genus Albericus by having a distinctly projecting snout comprising an extension of the nasal bones and the alary processes of the premaxillae. From other long-nosed Choerophryne it differs in the following characters: it is smaller (SUL 14.9 ���15.0 mm), than C. longirostris, C. microps, C. nigrescens and C. proboscidea (SUL> 15 mm), and further differs from these species in lacking an expanded disk on the first finger. Choerophryne epirrhina sp. nov. is larger than C. allisoni and C. burtoni (SUL Choerophryne epirrhina sp. nov is slightly larger than C. gracilirostris (SUL 14.9 ���15.0 vs. 13.5���14.7) and has a wider snout (IND /SUL 0.066���0.070 vs. 0.052���0.062). It is slightly larger than C. arndtorum (SUL 11.2���14.8) and has a longer snout (SL/SUL 0.23���0.24 vs. 0.19���0.22). Choerophryne epirrhina sp. nov. overlaps in size with C. rostellifer (male SUL 14.0��� 16.8 mm) and the geographically disjunct C. amomani (11.8���15.1 mm) and C. bryonopsis (15.0��� 17.9 mm). It differs from C. amomani and C. bryonopsis in lacking an expanded disk on the first finger, having a longer snout (OHG/SUL 0.09���0.10 vs. 0.06���0.07 and 0.05���0.06 respectively), and smaller eye diameter (ED/SUL 0.09���0.10 vs. 0.11���0.13 for both C. amomani and C. bryonopsis). Morphologically, this species is difficult to distinguish from C. rostellifer but it has a different call and is also genetically divergent (see below). Key diagnostic features of long-nosed Choerophryne found on the central cordillera of New Guinea are summarised in Table 2. In having calls normally containing four notes C. epirrhina sp. nov. differs from C. allisoni (5���6 notes), C. amomani (1���2 notes), C. bryonopsis (1 note), C. burtoni (5���6 notes), C. microps (5���9 notes), C. nigrescens (6���10 notes) and C. proboscidea (3 notes). Choerophryne arndtorum produces four-note calls, but the notes are shorter (0.13��� 0.18 s vs 0.20��� 0.25 s) and produced at a faster rate (2.7���3.4 /s vs. 2.0��� 2.3 /s). Choerophryne gracilirostris makes 3���5 -note calls with shorter notes (0.014���0.042 vs. 0.20��� 0.25 s) produced at a much faster rate (7.2���13.3 /s vs. 2.0��� 2.3 /s). Choerophryne longirostris and C. rostellifer also sometimes make four-note calls; but C. longirostris notes are much longer (0.45���0.52 vs. 0.20��� 0.25 s) and produced at a much slower rate (0.69 vs. 2.0��� 2.3 /s); C. rostellifer notes are much shorter (0.066���0.085 vs. 0.20��� 0.25 s) and produced at a much faster rate (~ 7 /s vs 2.0��� 2.3 /s). Description of holotype. Adult male with a short ventral incision on right side, liver removed, and measurements (in mm) of: SUL 14.9; TL 6.1; HW 5.1; HL 5.3; ED 1.5; END 2.5; IND 1.0; SL 3.4; OHG 1.4; F 1 D 0.4; F 3 D 0.7; T 1 D 0.3; T 4 D 0.8. Head rather narrow (HW/SUL 0.34), canthus rostralis weakly defined and smoothly rounded, loreal region very slightly concave. Snout narrow, acute and projecting well beyond anterior rictus of lower jaw (OHG 1.4 mm), tip rounded in dorsal and lateral profiles, ventral surface of tip with a broad, unpigmented medial ridge, clearly defined anteriorly and becoming shallow and indistinct towards anterior rictus of the upper jaw. Nares oval, narrowing slightly posteriorly, located close to tip of snout, oriented ventrolaterally and not visible in dorsal view; internarial distance moderately large (IN/SUL 0.066). Eye rather small (ED/SUL 0.099), transverse diameter less than length of snout, pupil horizontal. Tympanic annulus visible along the ventral and anterior edges of tympanic membrane, not visible dorsally or posteriorly. Urostyle projecting slightly beyond vent. Dorsum relatively smooth except for a very faint vertebral ridge extending from snout tip to urostyle tip; scattered tubercles across flanks, densest anteriorly; venter smooth. Fingers moderately long, with distinct circum-marginal grooves, relative lengths III>IV>II>I, finger I more than half length of II, fingers II���IV with distinctly expanded discs up to 1.6 times width of penultimate phalanx, finger I without disk and with only faint circum-marginal groove; subarticular tubercles, metacarpal tubercules and webbing absent. Legs moderately long (TL/SUL 0.41). Toes moderately wide, relative lengths IV>V���III>II>I, toe I just over half length of II, all toes except I with small terminal discs up to 1.2 times width of penultimate phalanx and with circum-marginal grooves; metatarsal tubercles and webbing absent. Toe discs wider than finger disks (F 3 / T 4 0.91). In preservative, ground colour of dorsum buff with dense dark grey-brown maculations, slightly denser anterior to the tympana, lending the head an overall darker appearance. Dorsum dominated by a longitudinally oriented, medium-brown, medially constricted blotch, wider posteriorly than anteriorly and with prominent but weakly defined darker brown edging. Inguinal region with a pair of indistinct dark-brown lumbar ocelli, each containing a distinct pale-cream medial spot. Vent bordered laterally and dorsally by a dark-brown arc, urostyle tip pale buff. Exposed dorsal and lateral surfaces of limbs pale buff, with extensive tan maculations and dark-brown blotches forming transverse bands at the midpoints of the thigh, shank, foot, forearm and hand, giving an overall barred appearance. Dorsal surfaces of hands and feet pale buff with extensive pale-brown flecking and darkerbrown blotches around the base of the distal phalanges. Venter and ventral surface of limbs pale buff with fine tan maculations. Palmar and plantar surfaces largely unpigmented except for scattered light-tan speckling. Variation. Measurements for the type series are given in Table 1. The paratype is of similar dimensions to the holotype, but has a slightly longer snout (OHG/SUL 0.10 vs. 0.094). Tuberculation of the flanks is sparser than on the holoype, and the slope of the loreal region is straight (vs. slightly concave). Like the holotype, the paratype also has a medium-brown dorsal blotch with darker-brown edging and distinct pale-cream inguinal spots, limbs with dark-brown transverse bands, and ventral colouration of pale buff with extensive tan speckling. Internal oral features of the holotype are not visible so those of the paratype are presented here: tongue oval, slightly bifurcated at tip; two curved prepharyngeal ridges present, vomerine teeth absent, vocal slits present slightly anterolateral to base of tongue. Appearance in life. Photographs in life (Figure 1 A) of the holotype taken at night show a greater degree of contrast between light and dark-brown dorsal and ventral areas than is evident in preservative. Dorsum grey, with dense brown and orange maculations (giving an overall brownish-grey appearance), overlain with darker-brown blotching and numerous smaller grey spots. Ventral surfaces tinged blue-grey, with scattered pale-grey blotches; belly and ventral surface of the snout overlain with extensive orange flecking; similar flecking is present but less dense on the throat and thighs. Iris black with small bronze flecks, pupil horizontal with a thin reddish-orange border. Dorsal surfaces of torso and legs with numerous pale-grey tubercles of heterogeneous size (less conspicuous in preservative). Advertisement call. Ten calls produced by the holotype were analysed, including nine consecutive calls produced in an uninterrupted series. A single complete call from the holotype is presented in Figure 2 A. The series of nine calls lasted 79.2 seconds and was produced at a rate of 0.1 calls/s. Energy was focused in two major frequency bands, one at around 3,000 hz and the other at around 3,800 hz. In seven of the ten calls analysed the dominant frequency was between 3692 and 3884 hz (mean = 3829 hz, SD = 71.4), while in the remaining three calls the dominant frequency was 2942���3037 hz (mean = 2999 hz, SD = 50.7). Calls contained 3���4 notes (nine of 10 calls had 4 notes) and were 1.13��� 1.70 s long (mean = 1.5, SD = 0.15), but this narrowed to 1.53��� 1.70 s (mean = 1.59, SD = 0.05) when the only 3 -note call was excluded from analysis. Note length was similar both within and between calls, ranging from 0.20��� 0.25 s (mean = 0.22, SD = 0.01; n = 39), and notes were produced at a rate of 2.0��� 2.3 /s (mean = 2.22, SD = 0.07; n = 10 calls). Notes contained 27���31 pulses produced at a rate of 117���141 pulses/s (mean = 127.8, SD = 6.51; n = 20 notes). Etymology. From the Greek, epirrhinos, meaning ���long-nosed���. Natural history and distribution. Choerophryne epirrhina sp. nov. is known from one location in primary foothill rainforest on the northern side of Papua New Guinea���s central cordillera at an altitude of 850 m. Males were calling after rain at night, from within curled leaves that were accumulated at the base of shrubs, within 20 cm of the ground. Given the extent of suitable habitat in the area, this species probably has a broad distribution along the northern slopes of the central cordillera. Choerophryne epirrhina sp. nov. and Choerophryne rostellifer are also very similar in external appearance, and some specimens from elsewhere on the central cordillera referred to the latter species (Kraus & Allison 2001) may also actually represent the former., Published as part of Iannella, Amy, Oliver, Paul & Richards, Stephen, 2015, Two new species of Choerophryne (Anura, Microhylidae) from the northern versant of Papua New Guinea's central cordillera, pp. 332-340 in Zootaxa 4058 (3) on pages 333-336, DOI: 10.11646/zootaxa.4058.3.2, http://zenodo.org/record/244825, {"references":["Kraus, F. & Allison, A. (2001) A review of the endemic New Guinea microhylid frog genus Choerophryne. Herpetologica, 57, 214 - 232."]}

Skipwith, Phillip L., Oliver, Paul M. (2014): A new Gehyra (Gekkonidae: Reptilia) from New Guinea with unique caudal scalation. Zootaxa 3827 (1): 57-66, DOI: http://dx.doi.org/10.11646/zootaxa.3827.1.5

Varanus hamersleyensis sp. nov. Southern Pilbara Rock Goanna (Figs. 8, 9, 10) Material examined. Holotype: WAM R 145733, male, Hamersley Range, Weeli Wolli Creek (22 º 52 ' 40 "S 119 º 14 ' 24 "E), Western Australia, Australia, R. Teale & G. Harold, 28 April 2003. Fixed in 10 % formalin, stored in 70 % ethanol, liver sample stored in – 80 ºC ultrafreezer at WAM. Paratypes: All from Western Australia. WAM R 125456 (male)— 30 km E Newman (23 º 19 'S 120 º02'E); WAM R 125766 (female)—Circular Pool, Dales Gorge (22 º 38 ' 36 "S 118 º 33 ' 47 "E); WAM R 129628 (male)— 120 km NW Newman (22 º 55 'S 119 º01'E); WAM R 140734 (female)—Rocklea Station (22 º 56 ' 55 "S 117 º 16 '0 7 "E); WAM R 164579 (male)— 8.5 km NE Mount Rica (21 º 57 '0 8 "S 116 º 29 ' 18 "E). Diagnosis and comparison. Differs from all Australian congeners by the following combination of characters: size moderately small (SVL up to 160 mm); slender build; tail long (between 168–207 % of SVL) and thin, circular in cross section at midpoint, midbody scales 120–122; dorsal and lateral scales weakly keeled, enlarged keeled ventrolateral scales on each side of and posterior to vent in 4–6 rows, scales on top of head smooth; supraoculars gradually merging with larger interoculars; dorsal scales elongate, ovate; nostrils high and oriented dorsolaterally, dorsal and lateral surfaces of body with subdued irregular dark brown to black spots or vermiculations on a dark reddish-brown background, small ocelli on the dorsal surface of the hindlimbs only, and tail without prominent dark bands. Varanus hamersleyensis sp. nov. differs from its sister species V. pilbarensis in its overall darker colouration, presence of small whitish ocelli usually on the dorsal surface of the limbs only (v. large greyish ocelli on the dorsal and lateral surfaces of body and limbs), largely unbanded tail (v. strongly banded tail) and smaller and less elongate dorsal scales (Fig. 3 B). Description of holotype (Fig. 8). Small-bodied Varanus with depressed body. Head elongate and depressed in lateral view; upper rostrum slightly protruding over lower jaw, genial groove distinct. Tail long and gradually tapering, moderately dorsoventrally depressed proximally, circular centrally, becoming more triangular distally. Eyes large with scaly upper and lower eyelids; closer to nostril than to ear opening; supraocular brow present above eyes. Snout moderately long, rounded in dorsal and lateral views; nostrils large and visible from above, wider than high, oriented dorsolaterally, approximately equidistant between tip of snout and anterior edge of eye. Ear openings much higher than wide, oriented distally at 45 º to the corner of jaw. Head scales small, smooth and unornamented; supraocular scales smaller than those on the rostrum, frontal and parietal regions; canthus rostralis weak; rostral scale twice as large as adjacent scales, almost rounded, slightly narrower anteriorly; mental scale higher than wide and almost rectangular. Limbs well-developed and moderately slender; ventral surface of digits with enlarged paired circular lamellae, scales on manus and pes otherwise consisting of conical tubercles; claws darkly pigmented, relatively short, very thick, laterally compressed and with sharp, recurved tips. Dorsal primary scales small, non-overlapping, relatively uniform and ovate, approximately twice as long as wide and with a small central keel; bordered by row of small granules; ventral scales non-overlapping and more rectangular, approximately 50 % longer and 100 % wider than dorsal scales, grading smaller on the throat and vent; caudal scales undifferentiated on dorsal and ventral surfaces, at base of tail similar in size to dorsal scales on torso, grading to larger with moderate keels in central section of tail and narrow with sharp keels towards tip; cloacal region with paired set of 4-6 curved lateroventral rows of long, narrow pointed scales, increasing in height, size and spinosity distally. In preservative, ground colour of dorsum dark brown, slightly lighter on head and neck. Patterning on head includes black spots on some individual scales and dark brown labial bars, dorsal surfaces of torso with mixture of irregular black spots on individual scales or spots coalesce to form narrow, wavy transverse lines. Light brown tail with irregular dark brown to black spots. Forelimbs light brown, hindlimbs darker brown with small whitish ocelli becoming more prominent distally. Ventral surfaces greyish. Variation. Summary mensural and meristic data for the type series of V. hamersleyensis sp. nov. is presented in Tables 1 and 3. In preservative, the ground colouration is always brown (Fig. 10). All specimens have irregular dark spots on some individual scales or vermiculations across the dorsum, although these vary in degree and intensity; on one individual (WAM R 125766) the entire dorsum has very fine spots with barely any indication of alignment, while at the other extreme (WAM R 140734) the dense spots align to form regular wavy lines. Most individuals have some dark spotting on the tail, although one individual (WAM R 164579) has almost none. The venter sometimes features relatively bold greyish bands that become more narrow and wavy on neck and throat. Two juvenile specimens (WAM R 69695: SVL 61.6 mm; WAM R 125100: 70 mm) are more brightly patterned than adults with much larger spots on dorsum, more contrasting and strongly defined ventral surface bands and obviously ringed tails. Colouration in life. The following description of colouration in life is based on Figs. 9 A, B and a photograph from Mount Channar near Paraburdoo (Ehmann 1992). Dorsal ground colour of head, torso and limbs dark reddishbrown, often tending slightly paler on head. Head and neck with irregular subdued small blackish-brown to black spots, snout forward of nostrils often dark brown, sides of head with yellowish tinge. Labial scales with blackishbrown vertical bars. Dorsal surface of torso with numerous small blackish-brown to black spots, often coalescing into narrow (one scale wide) transverse bars or wavy lines and forming a fine black reticulum. Forelimbs reddish brown with dark flecks, hindlimbs noticeably darker brown with small whitish ocelli. Digits on both limbs with indistinct thin pale yellowish bands. Ground colour of the tail light brown, extensively overlain with darker-brown to black spots along its length. Ventral surface of the throat, neck, limbs and body greyish-white. Habitat and ecology. Varanus hamersleyensis sp. nov. is strictly saxicolous and closely associated with rocky habitats such as banded ironstone rock faces and gorges (Fig. 11). It shelters in overhangs, crevices, cavities and occasionally under exfoliated rock slabs. Typical vegetation on these faces varies, but often includes Eucalyptus and Triodia (Spinifex). Field observations indicate a preference for flat, horizontal sheet ironstone with an eastwest aspect to allow optimum basking in the morning. It is extremely swift and agile; ascending vertical rock faces with ease, and quickly takes shelter in deep crevices when disturbed. As with V. pilbarensis (Ehmann 1992; B. Maryan pers. obs.), this species has been observed active after sunset on warm, humid nights. Field observations indicated that activity periods vary seasonally, peaking during the hotter and humid months from October–April and declining when conditions are dry from May–September. The sex ratio among the type series of V. hamersleyensis sp. nov., including the misidentified paratype of V. pilbarensis (WAM R 14901) is skewed (5 adult males and 2 adult females), a bias which seems to be consistent with museum series of other opportunistically collected varanids (Greer 1989) and may reflect different activity periods of the sexes. During surveys in the eastern Hamersley Range, most individuals captured were also males (B. Maryan and M. O’Connell, unpublished data). Distribution. Known only from the Hamersley Range, a heavily dissected region of banded ironstone occupying the southern part of the Pilbara craton of Western Australia. Records span the Hamersley Range for over 300 km from east of Newman (WAM R 125100, WAM R 125456) west to Mount Rica (WAM R 164579) and south to Rocklea Station (WAM R 140734) (Fig. 2). Etymology. Named for the Hamersley Range of Western Australia, the region to which it is restricted. This is the most prominent mountainous area in Western Australia (including Mount Meharry, 1245 m, and the highest peak in the state) and home to a suite of endemic taxa. Used as a noun in apposition. Remarks. This species appears to be secure and widespread throughout the Hamersley Range, and occurs in popular tourist destinations such as at Karijini National Park (one of the largest National Parks in Western Australia).

Maryan, Brad, Oliver, Paul M., Fitch, Alison J., O'Connell, Morgan (2014): Molecular and morphological assessment of Varanus pilbarensis (Squamata: Varanidae), with a description of a new species from the southern Pilbara, Western Australia. Zootaxa 3768 (2): 139-158, DOI: http://dx.doi.org/10.11646/zootaxa.3768.2.3

Varanus pilbarensis Storr, 1980 Northern Pilbara Rock Goanna (Figs. 4, 5, 6) Material examined. Holotype: WAM R 39782, male, Chichester Range, Cockeraga River (22 ��03'S 118 �� 48 'E), Western Australia, Australia, J.C. Wombey, 7 April 1971. Paratypes: All from Western Australia. WAM R 10811 (male)���Abydos (21 �� 25 'S 118 �� 55 'E); WAM R 13082 * (unsexed)���Woodstock Station (21 �� 37 'S 118 �� 57 'E); WAM R 14901 ** (male)���Dales Gorge, Hamersley Range (22 �� 30 'S 118 �� 36 'E); WAM R 20017 (male)���Black Hill Pool (21 �� 19 'S 117 �� 15 'E); WAM R 28011 ��� 12 (males)���Woodstock Station (21 �� 37 'S 118 �� 57 'E); WAM R 39158 (male)��� 2 km W Nullagine (21 �� 54 'S 120 ��05'E); WAM R 60432 (female)���Woodstock Station (21 �� 37 'S 118 �� 57 'E). *Poor specimen not included in morphological examination. ** Represents new taxon described below. Extended diagnosis and comparison. Differs from all Australian congeners by the following combination of characters: size moderately small (SVL up to 180 mm); slender build; tail long (between 170���211 % of SVL) and thin, circular in cross section at midpoint, midbody scales 118���128; dorsal and lateral primary scales weakly keeled, enlarged keeled ventrolateral scales on each side of and posterior to vent in 4���6 rows, scales on top of head smooth; supraoculars gradually merging with larger interoculars; dorsal scales elongate, ovate; nostrils high and oriented dorsolaterally, dorsal and lateral surfaces of body with distinct transverse series of large ocelli on a light reddish-brown background, and tail with prominent alternating light and dark bands. Varanus pilbarensis differs from its sister species in the Hamersley Range (described below) in its overall paler colouration, large greyish ocelli on the dorsal and lateral surfaces of body and limbs (v. small whitish ocelli on limbs only), strongly banded tail (v. completely or mostly unbanded) and larger and more elongate dorsal scales (Fig. 3 A). Re-description of holotype (Fig. 4). Small-bodied Varanus with a depressed body. Head elongate and depressed in lateral view; upper rostrum slightly protruding over lower jaw, genial groove distinct. Tail long and gradually tapering, moderately dorsoventrally depressed proximally, circular centrally, becoming more triangular distally. Eyes large with scaly upper and lower eyelids, closer to nostril than to ear opening; supraocular brow present above eyes. Snout moderately long, rounded in dorsal and lateral views; nostrils large and visible from above, oriented dorsolaterally, approximately equidistant between tip of snout and anterior edge of eye. Ear openings much higher than wide, oriented distally at 45 �� to the corner of jaw. Head scales small, smooth, supraoculars gradually merging with the larger interoculars; canthus rostralis weak (not entirely absent as stated by Storr 1980), demarcated by low ridge of larger scales anterior to eye and contacting nostril; rostral and mental scales higher than wide, twice the size of adjacent scales. Limbs well-developed and moderately slender, ventral surface of digits with enlarged paired circular lamellae, scales on manus and pes otherwise consisting of conical tubercles; claws short and thick, with strongly recurved tips. Dorsal primary scales small, relatively uniform and ovate with small central keel, bordered by row of small granules; ventral scales larger and much longer than wide; caudal scales undifferentiated on dorsal and ventral surfaces, at base of tail similar in size to dorsal scales on torso, grading to larger with moderate keels in central section of tail and narrow with sharp keels towards tip; cloacal region with paired set of 4���6 curved ventrolateral rows of long, narrow pointed scales, increasing in height, size and spinosity distally. In preservative, ground colour light brown, with reddish tinge on head and limbs, and tending greyer on flanks. Patterning on head includes light brown spotting and similarly coloured bars on labials. Dorsal surfaces and base of tail with dark brown spots (prominent on tail) coalescing to form narrow, wavy transverse lines. Limbs with light grey ocelli, larger and more prominent on hindlimbs. Dark brown transverse bands present from mid to distal point of tail. Ventral surface white with faint light grey bands. Variation. Tables 1���2 summarises the mensural and meristic data for V. pilbarensis. In preservative, dorsal patterning in most individuals is similar (Fig. 6), consisting of fine dark brown spots on a lighter brown background. In some individuals (WAM R 100766, WAM R 165513) the entire dorsum has widely scattered spots on individual scales, while in others (WAM R 102103, WAM R 163916, WAM R 170132) the spots coalesce to form irregular transverse lines. The smallest individual (WAM R 132659) is very boldly marked with dorsal spots almost forming wavy bars across dorsum. One individual (WAM R 110941) has a uniform dark brown body and lighter head with very indistinct ocelli on the dorsum and a strongly banded tail. All specimens have strongly banded tails. The large, pale grey ocelli are faint on some individuals (WAM R 100766). V. pilbarensis V. hamersleyensis sp. nov. Colouration in life. The following description of colouration in life is based on Figs. 5 A, B and field observations of V. pilbarensis from the Chichester and surrounding northern ranges. Dorsal ground colour of head, body and limbs light reddish-brown, slightly paler (tending towards yellowish) on head. Dorsal surfaces of head and neck distinctly spotted or flecked with dark reddish-brown, sometimes coalescing to form narrow wavy, transverse bands. Distinct reddish-brown to black vertical bars on labial and temporal regions. Upper surface of dorsum and base of tail with numerous small blackish-brown to black spots that coalesce into narrow (one scale wide) transverse bars or wavy lines. Between these spots are transverse rows of large greyish ocelli. These ocelli vary in degrees of intensity and with body size but are clearly discernible posteriorly at any stage of maturity. Limbs dorsally and laterally with irregular pale grey or whitish ocelli (often dark-edged and/or dark-centred) and, larger and brighter on hindlimbs. Ground colour of proximal section of tail light brown or greyish, often with large greyish ocelli; ground colour of distal section of tail grading from cream to white with dark reddish-brown to black rings, giving a distinct banded appearance. Ventral surfaces of throat, neck, limbs and body greyish-white, often with irregular transverse grey bands or reticulations. Juveniles are more simply and very brightly patterned, with large black-edged pale ocelli on dorsum and limbs, distinct reddish-brown heads and black and white banded tails. Habitat. This species is closely associated with rocky habitats (Fig. 7), typically undulating granite and greenstone terrain with significant areas of basaltic ranges (tumbled boulder piles are prominent) and dissected plateau, vegetated with Triodia, sparse shrubs and eucalypts (Mckenzie et al. 2009). Distribution. Endemic to the northern Pilbara region of Western Australia, largely to the north of the Fortescue River Basin and centred on the Chichester Range. Records extend from the Burrup Peninsula and West Intercourse Island in the northwest, east to the Ord Ranges, Yarrie Mine and Carawine Gorge. There is a single record south of the Fortescue River in the western Chichester Range near Pannawonica (Fig. 2). Etymology. Named for the Pilbara region of Western Australia, the region to which it is restricted. Used as a noun in apposition. Remarks. This species appears to be secure and widespread throughout the Chichester Range and other northern ranges, and occurs in popular tourist destinations such as at Millstream Chichester National Park. This species can be locally abundant in suitable rocky areas. Varanus pilbarensis is illustrated in numerous publications (e.g. Storr et al. 1983: Plate 15; Vincent & Wilson 1999: 75; Pianka et al. 2004: 430; Eidenm��ller 2007: 105; Wilson & Swan 2013: 429)., Published as part of Maryan, Brad, Oliver, Paul M., Fitch, Alison J. & O'Connell, Morgan, 2014, Molecular and morphological assessment of Varanus pilbarensis (Squamata: Varanidae), with a description of a new species from the southern Pilbara, Western Australia, pp. 139-158 in Zootaxa 3768 (2) on pages 145-149, DOI: 10.11646/zootaxa.3768.2.3, http://zenodo.org/record/252057, {"references":["Storr, G. M. (1980) The monitor lizards (Genus Varanus Merrem, 1820) of Western Australia. Records of the Western Australian Museum, 8, 237 - 293.","Storr, G. M., Smith, L. A. & Johnstone, R. E. (1983) Lizards of Western Australia. II. Dragons and Monitors. Western Australian Museum, Perth, Western Australia, Australia, 113 pp.","Vincent, M. & Wilson, S. (1999) Australian Goannas. New Holland Publishers, Sydney, New South Wales, Australia, 152 pp.","Pianka, E. R., King, D. R. & King, R. A. (2004) Varanoid Lizards of the World. Indiana University Press, Bloomington, Indiana, U. S. A., 588 pp.","Eidenmuller, B. (2007) Monitor Lizards, Natural History, Captive Care & Breeding. Revised, enlarged and updated edition. Edition Chimaira, Frankfurt am Main, Germany, 176 pp.","Wilson, S. & Swan, G. (2013) A Complete Guide to Reptiles of Australia. Fourth Edition. New Holland Publishers, Sydney, New South Wales, Australia, 592 pp."]}

Gehyra serraticauda sp. nov. (Figs 1���3) Holotype. MCZ R 7314, adult male, purchased from Antwerp Edgar Pratt and with collection locality recorded as Fakfak, Onin Peninsula (~ 2 �� 55 '"S, 132 �� 18 'E), West Papua Province, Indonesia. Date of original collection unknown, but lodged in the MCZ collection in 1909. Additional material. Uncollected specimen, from near Kamaka (formerly Warika) Village, 45 km SSE of Kaimana, Triton Bay, (03�� 46 ��� 14 ���S, 134 �� 10 ��� 14 ���E), West Papua Province, Indonesia, 150���160 m a.s.l., collected, photographed and released by Dmitry Telnov, 10 September 2010. Diagnosis. Gehyra serraticauda is distinguished from other Gehyra species by the following suite of characters: moderately large size (91 mm SVL), prominent popliteal fold on the hindlimbs, high number of digital lamellae (finger IV = 16, toe IV = 17), distal lamellae deeply notched, rostral concave, supranasals small and widely separated by numerous small internasals, preanal pores arranged in a long single continuous chevron (36), and original tail strongly compressed dorsoventrally and adorned with a continuous series of acuminate scales on the lateral edges. Comparisons. A summary of comparative data for Melanesian Gehyra is given in Table 1. The combination of moderately large body size (> 90 mm) and distinctive acuminate lateral scales on the tail distinguish this species from all other Gehyra. Gehyra serraticauda sp. nov. specifically differs from Gehyra mutilata, G. papuana and the types of G. lampei and G. interstitialis in having very distinct lateral caudal serrations (versus minute or none), a much larger adult size (> 90 mm versus versus G. interstitialis (actual types lost), Gehyra serraticauda sp. nov. further differs in having a notched rostral (versus quadrangular) and U-shaped mental (versus triangular). Gehyra leopoldi is a poorly known taxon that may be synonymous with G. mutilata (Bauer & Henle 1994); it is distinguished from G. serraticauda sp. nov. by its much smaller size (44 mm), lower number of subdigital lamallae (always versus usually> 9 (exceptions being finger I and toe I)), and presence of only a single internasal in a dorsal concavity of the rostral. Gehyra serraticauda sp. nov. is similar in size to G. baliola, G. barea and G. oceanica, but again differs in having well developed tail serrations. It is further differentiated from G. oceanica by having deeply notched lamellae (versus shallowly notched and undivided), the presence of numerous small internasal scales (versus absent), enlarged subcaudals (versus small), and the presence of a prominent popliteal skinfold (versus absent). Gehyra serraticauda sp. nov. can be distinguished from a final large species of Papuan Gehyra, G. membranacrularis by its deeply notched subdigital lamellae (versus shallowly notched and undivided) and smaller and more numerous internasals (~ 10 versus 1���4). Gehyra serraticauda sp. nov. differs from Gehyra marginata Boulenger from the Maluku Islands of Indonesia (just west of New Guinea) in its smaller adult size (versus > 130 mm), divided digital lamellae (versus undivided), lower number of lamellae (digit IV manus = 17 versus 20���25, digit IV pes = 17 versus 20���23), by the presence of many small internasals (versus a single large internasal), well developed popliteal skin folds only (versus well developed lateral skin folds on the trunk and both antecubital and popliteal skin folds), and having enlarged polygonal subcaudals (versus small and relatively uniform). Two other gekkonid genera occuring in the Pacific region may also have flattened tails with lateral serrations. The three species of Perochirus from Micronesia and Vanuatu have dorso-ventrally flattened tails with very distinct lateral spines; but differ from all Gehyra in the presence of a well developed claw and free phalanx on the inner toe (versus vestigal or absent), and further differ from G. serraticauda sp. nov. specifically in lacking popliteal folds and having much more widely spaced lateral spines (one per tail segment) (Zug 2013). Hemidactylus includes a number of species that have tail serrations and expanded digital pads, but can again be readily distinguished by the presence of a distinct claw on all digits (greatly reduced on the inner toe in Gehyra serraticauda sp. nov.), and are also generally somewhat smaller than G.serraticauda sp. nov. Description of holotype.��� Adult male with expressed postcloacal pores and large flap of loosely attached skin on the right side of the head (Figs. 1 a���b): SVL 91.0 mm; TrK 46.3 mm; HW 17.1 mm; HL 21.8 mm; HD 10.8 mm; EN 7.6; IORB 8.6 mm; POM 2.9 mm; FA 12.3 mm; CS 13.3 mm; EYE 5.8 mm; EAR 2.1 mm; TL(total) 83.0; TL(original) 65.0; IN 10; INT 5; SUPR 13; INFR 11; LAMF 4 17; LAMT 4 17. Head triangular, longer than wide (HW/HL= 0.78) moderately large (HL/SVL = 0.24) and deep (HD/HL = 0.50); anterior left dentary fractured, posterior corner of jaw upturned. Rostrum long and robust (EN/HL = 0.35) with distinct dorsal concavity, transverse fold of skin extends across tip of rostrum. Rostral with deep dorsal notch, in contact with two supranasals and five small internasals along dorsal edge (Fig. 2 c). Supranasals separated by a high number of small internasals (n = 10) and up to five in transverse series. Nares contacting one supralabial, rostral, one large supranasal and one large postnasal; supralabials 11���12; infralabials 12. Mental U-shaped, bordered by oblong post-mentals. Pupil partially dilated, somewhat elliptical with smooth margins and limited crenulations (Fig. 2 d). Body long and robust (TrK /SVL = 0.40), thorax flexed prominently to the left. Skin on dorsum and venter smooth and composed entirely of small, flat, granular scales. Limbs relatively short and stout. Digits on both the fore and hind limbs with prominent and expanded pads (finger pads 1.1���1.6 times minimum width of finger, toe pads 1.2���1.5 minimum width of toe) (Fig. 2 a���b); penultimate phalanx free and well developed on all digits except finger I and toe I. The scansorial pad of digit I of both manus and pes is narrower relative to length than the other digits. Distal lamallae (excluding penultimate lamallae) deeply notched on fingers and toes, lamellar counts for all digits (total/deeply notched) as follows: fingers I = 11 / 4, II = 12 / 5, III = 16 / 6, IV = 17 / 7, V = 16 / 7; toes I = 14 / 6, II = 16 / 7, III = 17 / 8, IV = 17 / 8, V = 16 / 7. Basal webbing between digits limited, never reaches first phalangeal joint. Precloacal and femoral pores (n = 36) arranged in a single curved row terminating halfway along the femur (Fig. 2 f). Hemipenal bulge present but not pronounced, single row of three short cloacal spurs angled posteriorly on each bulge. Tail strongly compressed dorsoventrally, approximately 1.5 times wider than high, 83.0 mm in total length (65.0 mm original, 18.0 mm regenerated). Original portion partially autotomised 24.0 mm from the vent, with a distinct medial groove on the ventral surface, a single row of greatly enlarged pentagonal subcaudals extending its full length, and a continuous lateral fringe of densely packed acuminate scales extending from just posterior to the hindlimbs to the end of the original tail (Fig. 2 e). Scalation on the regenerated section of the tail is substantially smaller and more irregular and heterogeneous than that of the original, although the subcaudals are still relatively enlarged. In preservative, dorsum beige with irregular dark grey patches on the left shoulder, lateral regions of torso, pelvic area, and distal portion of the tail. Dorsal surface of the hands and feet beige like the dorsum of the body but with a slight reddish tinge, giving an overall darker appearance; lamellae of all digits beige becoming slightly darker distally. Colouration on the regenerated tail light reddish-grey. Variation. Photographs of the uncollected specimen from near Kamaka Village kindly provided to us by Dmitry Telnov (Fig. 3 a���c), specimen details above) show that it has a deeply divided rostral, prominent popliteal folds and acuminate scales on the lateral edges of the original tail. It is on the basis of these characters and its large size (field measured SVL ~ 120mm) that we assign this individual to the G. serraticauda sp. nov. When originally captured the colouration of this specimen was as follows; dorsum silvery grey with very extensive terracotta mottling and numerous indistinctly edged transverse bands along the dorsum; fore and hindlimbs predominently terracotta with no clear pattern but some silvery flecking; head with similar mottled colouration to the body and distinct silvery loreal and postorbital stripes; regrown tail silvery brown with no clear pattern; supralabials, infralabials and region of venter visible in photographs yellow; iris light olive green and pupil ellipitical with scalloped edges. In photos taken at a later time prior to release the dorsal colouration is greatly faded and mostly silvery grey with a brownish tinge towards the anterior regions and the only clear patterning being a small number of silvery spots and stripes on the head and neck. Distribution and ecology. Presuming the collection data for the holotype is accurate; this distribution of this species extends from Fakfak on the Onin Peninsula east as far as the Triton Bay region. Collection information for the holotype indicates that it was purchased from local people, and it thus seems likely that it is from a locality that is within walking distance of Fakfak. However, more fieldwork is required to confirm that this species occurs in this area. The Triton Bay specimen was collected in primary lowland rainforest on limestone in the environs of Kamaka Village. It was found during the day while hiding under bark on a dead standing tree. Like many Gehyra the strong dorsal patterning on this specimen varied over short periods of time (King & Horner 1989). Nothing is known about the collection locality and habitat of the holotype. Eytmology. The specific epithet is a feminine combination of the Latin adjective ���serratus��� (notched like a saw) and the noun ���cauda��� (tail), and refers to the distinctive enlarged lateral scales on the tail of this species. Remarks. The taxonomic status and distribution of many Papuan Gehyra remains unclear (a situation exacerbated by low samples sizes for non-human commensal species and the loss and destruction of key types). New material and a proper phylogenetic analysis is required before G. serraticauda sp. nov. can be confidently placed in the phylogeny of Gehyra. However, it is superficially most similar to G. baliola and G. b a re a in overall size and proportions, the presence of a high number of scales between the nasals, and deeply notched lamellae. The distribution of this species also sits between the known range of G. baliola (southern New Guinea) and G. barea (probably widespread over islands just to the west of New Guinea), suggesting possible geographic turnover of ecologically similar and related taxa. Gehyra, for the most part, is a morphologically conserved taxon and the prominent lateral fringe of acuminate scales on the tail of G. serraticauda sp. nov. is quite unique (although Gehyra mutilata does have a similar, but much less prominent fringe). Many other gekkonids have flattened and or ornamented tails; ranging from prominent lappets (e.g., Ptyhcozoon sp.) to spiniform scales (e.g., Kolekanos plumicaudus and Phelsuma serraticauda). These structures may serve an array of functions, but are most frequently and most easily correlated with outline disruption and camouflage (Young et al. 2002; Heinicke et al. 2012)., Published as part of Skipwith, Phillip L. & Oliver, Paul M., 2014, A new Gehyra (Gekkonidae: Reptilia) from New Guinea with unique caudal scalation, pp. 57-66 in Zootaxa 3827 (1) on pages 58-63, DOI: 10.11646/zootaxa.3827.1.5, http://zenodo.org/record/286487, {"references":["Bauer, A. M. & Henle, K. (1994) Familia Gekkonidae (Reptilia, Sauria). Part 1. Australia and Oceania. Das Tierreich 109, 309 pp.","Zug, G. R. (2013) Reptiles and Amphibians of the Pacific Islands: A Comprehensive Guide. University of California Press, Berkeley, California, 306 pp.","King, M. & Horner, P. (1989) Karyotypic evolution in Gehyra (Gekkonidae: Reptilia). V. A new species from Papua New Guinea and the distribution and morphometrics of Gehyra oceanica (Lesson). Beagle, 6, 169 - 178.","Young, B. A., Lee, C. E. & Daly, K. M. (2002) On a flap and a foot: aerial locomotion in the '' flying' ' gecko, Ptychozoon kuhli. Journal of Herpetology, 36, 412 - 418. http: // dx. doi. org / 10.2307 / 1566185","Heinicke, M. P., Greenbaum, E., Jackman, T. R. & Bauer, A. M. (2012) Evolution of gliding in Southeast Asian geckos and other vertebrates is temporally congruent with dipterocarp forest development. Biology Letters, 8, 994 - 997. http: // dx. doi. org / 10.1098 / rsbl. 2012.0648"]}

Varanus Merrem, 1820 Type species: Lacerta varia Shaw in: White, 1790 (= Varanus varius, by subsequent designation of Gray, 1827). Diagnosis. The monotypic genus Varanus differs from all other squamate genera in possessing the following combination of character states: head long and flat, neck long; tongue long and forked, legs well developed, each with 5 strongly-clawed digits; tail long, gradually tapering, in cross section circular, rounded-triangular or laterally compressed; skin smooth to rough, somewhat loose-fitting; head covered by small juxtaposed polygonal scales and body and limbs by oblong or squarish, juxtaposed, non-glossy scales often bearing a low central keel., Published as part of Maryan, Brad, Oliver, Paul M., Fitch, Alison J. & O'Connell, Morgan, 2014, Molecular and morphological assessment of Varanus pilbarensis (Squamata: Varanidae), with a description of a new species from the southern Pilbara, Western Australia, pp. 139-158 in Zootaxa 3768 (2) on page 144, DOI: 10.11646/zootaxa.3768.2.3, http://zenodo.org/record/252057, {"references":["Merrem, B. (1820) Versuch eines Systems Amphibien. Tentamen Systematics Amphibiorum. Krieger, Marburg, 199 pp.","White, J. (1790) Journal of a voyage to New South Wales, with sixty-five plates of non descript animals, birds, lizards, serpents, curious cones of trees and other natural productions. Debrett, London, 299 pp."]}

Richards, Amy Iannella Stephen, Oliver, Paul (2014): A new species of Choerophryne (Anura, Microhylidae) from the central cordillera of Papua New Guinea. Zootaxa 3753 (5): 483-493, DOI: http://dx.doi.org/10.11646/zootaxa.3753.5.6