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+
+
+
+A new species of montane Prasinohaema (Sphenomorphini: Scincidae) from the South-fold Mountains of New Guinea
+
+
+
+Author
+
+Slavenko, Alex
+Cesar Australia, Brunswick, Victoria 3056, Australia
+
+
+
+Author
+
+Shea, Glenn M.
+Sydney School of Veterinary Science, B 01, University of Sydney, New South Wales 2006, Australia & Australian Museum Research Institute, Australian Museum, 10 William St, Sydney, New South Wales 2010, Australia & Herpetology Department, South Australian Museum, North Terrace, Adelaide, South Australia 5000, Australia
+
+
+
+Author
+
+Oliver, Paul M.
+School of Environment and Science, Griffith University, Nathan, Queensland, Australia, and Biodiversity and Geosciences, Queensland Museum, South Brisbane, Queensland 3300, Australia
+
+text
+
+
+Zootaxa
+
+
+2025
+
+2025-04-15
+
+
+5621
+
+
+2
+
+
+211
+230
+
+
+
+
+https://doi.org/10.11646/zootaxa.5621.2.3
+
+journal article
+10.11646/zootaxa.5621.2.3
+1175-5326
+15262507
+B85E0A5A-E44B-4752-B7FF-26202D039517
+
+
+
+
+
+
+
+Prasinohaema scurrula
+
+sp. nov.
+
+
+
+
+
+
+
+urn:lsid:zoobank.org:act:
+8BE96E88-5CFE-41EB-9908-68B7769FC4D6
+
+
+
+Figs. 2–6
+
+
+
+
+
+
+Holotype
+
+SAMA
+R65176
+(Field number SJR12137),
+Apalu Reke
+(
+-5.4862
+,
+142.3019
+),
+
+2,875 m
+a.s.l.
+
+,
+Muller Range
+,
+Hela Province
+,
+Papua New Guinea
+, collected by
+Stephen Richards
+on
+
+21 September 2009
+
+.
+
+
+
+
+
+Paratype
+.
+
+PNGNM 24992
+Tari Gap
+(nearest town Tari),
+
+2,700 m
+a.s.l.
+
+,
+Hela Province
+,
+Papua New Guinea
+, collected by
+Clifford Frith
+between
+
+24 December 1988
+
+to
+
+30 January 1989
+
+. Approximate coordinates are
+-5.9515
+,
+143.1463
+, georeferenced based on aerial photo in
+Frith & Frith (1990)
+, detailing the location of a
+
+Pteridophora alberti
+Meyer, 1894
+
+nest next to which the paratype was collected (C. Frith pers. comm.)
+
+.
+
+
+
+
+Diagnosis.
+
+Prasinohaema scurrula
+
+
+sp. nov.
+
+can be differentiated from all other
+
+Lobulia
+
+group skinks by the combination of adult SVL up to at least
+70 mm
+, snout short and relatively blunt; prefrontals not in contact, primary temporal not divided, parietals fully separated by interparietal, lower eyelid with two enlarged opaque scales centrally, more than two chin shields in medial contact, posterior chin shields separated from infralabials by a short row of 2–3 genials, subdigital lamellae only moderately expanded (distal lamellae typically around double width of proximal lamellae), lining of mouth blue in life, dorsal pattern of dark-brown crossbands, and venter with dark brown chequerboard pattern.
+
+
+
+
+Description.
+Snout short, dorsal margin convex in lateral profile; rostral wider than tall, in contact posteriorly with first supralabial, nasals and frontonasal, median lobe slightly rounded; frontonasal wider than long, in contact posteriorly with prefrontals and frontal, suture with frontal narrower than that with rostral and narrower than width of frontal at level of suture between second supraocular and frontoparietal; prefrontals moderately separated, in contact posteriorly with frontal, first supraocular and first supraciliary; frontal kite-shaped, longer than wide (
+paratype
+with anterior angle hived off as a small azygous scale, and posterior angle partially cleft longitudinally); one pair of frontoparietals, left overlapping right; supraoculars four, second extending most medially, third narrowest, first two contacting frontal, last three contacting frontoparietals; interparietal kite-shaped, shorter and narrower than frontal (
+paratype
+with anterior lobe of interparietal hived off as an azygous scale between frontoparietals), posteriorly completely separating parietals (
+Fig. 2
+); parietal eye spot absent (
+Fig. 3A&F
+).
+
+
+Nasal oblique, anteroventral to posterodorsal length shorter than anterodorsal to posteroventral length, contacting frontonasal dorsally and anterior loreal posteriorly; nostril central in nasal; a weak postnarial groove beginning ventral to nostril, arcing around posterior margin to posterodorsal side; no supranasal or postnasal; anterior loreal taller than long, partially (
+holotype
+) or completely (
+paratype
+) divided into smaller upper scale and larger lower scale, and dorsally contacting frontonasal and prefrontal, posteriorly contacting posterior loreal; posterior loreal squarish, but narrower ventrally, contacting prefrontal and first supraciliary dorsally, and first preocular and first presubocular posteriorly; on right side of
+paratype
+, posterior loreal divided into smaller upper and larger lower scale, together slightly taller than wide; supraciliaries nine (eight on R side of
+paratype
+), first four (three on R side of
+paratype
+) contacting first supraocular (fourth only narrowly so), first longer than the rest, narrowly separated from frontal, last two progressively arcing medially behind fourth supraocular, last overlapping parietal; a large anterior preocular wedged between first supraciliary and first presubocular, reaching posterior loreal anteriorly, followed by small second preocular that subtends both upper and lower palpebral rows; presuboculars three; postsuboculars four, first (lowermost) interdigitating between fifth and sixth supralabials, uppermost overlapped by last supraciliary, overlapping corner of parietal; upper palpebrals in contact with supraciliary row along most of its length, anterior part of row separated from supraciliaries by one (
+holotype
+) or 2L/3R (
+paratype
+) oblique scales; lower palpebrals shallow; the two central subpalpebrals enlarged, equivalent to a “window” but opaque; primary temporal single, overlapped by upper three postsuboculars and penultimate supralabial; secondary temporals two, upper longer than tall, overlapped by parietal, uppermost postsubocular and primary temporal; lower secondary temporal overlapped by upper secondary, primary and last supralabial; supralabials seven, first four squarish, last two pentagonal, first dorsally contacting nasal and narrowly anterior loreal (
+holotype
+) or just nasal (
+paratype
+), second contacting anterior loreal broadly and posterior loreal more narrowly, without (
+holotype
+) or narrowly with (
+paratype
+) contact with nasal as well, third narrowly contacting posterior loreal, broadly contacting first presubocular, and very narrowly contacting second presubocular, fourth contacting second and third presuboculars, fifth below centre of eye, separating pre- and postsubocular series, contacting 6L/5R (
+holotype
+) or 3L/2R (
+paratype
+) granules of lower eyelid; postsupralabials two; nuchals irregularly and asymmetrically differentiated,
+holotype
+with first pair of enlarged nuchals extending laterally to reach upper secondary temporals followed by 2L/1R narrower nuchals,
+paratype
+with two scales between paravertebral scales and upper secondary temporals bilaterally, and first few paravertebrals of irregular width; ear ovoid, taller than long, height about equal to height of lower eyelid, with two (
+holotype
+) or three (
+paratype
+) low lobules along anterior margin.
+
+
+
+FIGURE 3.
+
+Prasinohaema scurrula
+
+
+sp. nov.
+
+holotype (SAMA R65176). A) Dorsal view; B) Ventral view; C) Left front foot (ventral view); D) Left hind foot (ventral view); E) Head (left lateral view); F) Head (dorsal view); G) Head (ventral view).
+
+
+
+
+FIGURE 4.
+
+Prasinohaema scurrula
+
+
+sp. nov.
+
+paratype (PNGNM 24992). A) Dorsal view; B) Ventral view.
+
+
+
+
+FIGURE 5.
+A)
+
+Prasinohaema scurrula
+
+
+sp. nov.
+
+from Gigira Ridge, specimen not collected. B) Close up of the mouth of the Gigira Ridge specimen, showing the distinctive blue mouth lining. C)
+
+Prasinohaema scurrula
+
+
+sp. nov.
+
+holotype (SAMA R65176). D) & E) The type locality at Apalu Reke, Hela Province. F) Habitat at Tari Gap ~2,700 m a.s.l., approximate location of the collection locality of the paratype. All photographs by SJR.
+
+
+
+
+FIGURE 6.
+Map of New Guinea, with elevation above sea level in greyscale. Localities for examined material for different species of
+
+Prasinohaema
+sensu stricto
+
+(see text) are marked by different colours. Holotypes are marked by squares, paratypes by triangles, and photos by diamonds. Half-filled circles represent more than one species sampled at that locality. The coordinates for the holotype of
+
+P. parkeri
+
+are approximate.
+
+
+
+Mental similar width to rostral, but shallower on midline; postmental slightly narrower than mental; infralabials seven (
+holotype
+) or 8L/7
+R
+(
+paratype
+), first two contacting postmental, second to third (L on
+holotype
+,
+R
+on
+paratype
+) or second to fourth contacting chin shields, remainder separated by one or more rows of intervening scales;
+paratype
+with first two pairs of chin shields in median contact, third pair of chin shields medially separated by two scales; fourth pair of chin shields separated medially by three scales;
+holotype
+with asymmetrical chin shields, first two on left contacting first on right, second on right narrowly separated from its partner on the left by a single scale, third pair separated by two scales, but right scale laterally shortened to allow contact between second and fourth chin shield of that side, fourth pair separated by three/four irregular scales.
+
+
+Body scales glossy, in 32 (
+holotype
+) or 34 (
+paratype
+) longitudinal rows at midbody; dorsals irregularly longitudinally grooved (
+holotype
+) or smooth (
+paratype
+), slightly wider than long, paravertebral series not enlarged, 62 to level of anterior margin of hindlimb (both specimens), 65 (
+holotype
+) or 66 (
+paratype
+) to posterior margin of hindlimb; medial precloacals enlarged and laterally overlapping the adjacent precloacals; median subcaudals slightly wider than laterally adjoining subcaudals; tip of tail rounded with slight development of a ventral “pad”.
+
+
+Limbs pentadactyle, third and fourth fingers of similar length; fourth toe longest. On toes, basal subdigital lamellae only slightly wider than distal subdigital lamellae, and width gradually diminishes distally.Fourth toe slightly dorsally arched at second interphalangeal joint; fifth toes slightly arched at first interphalangeal joint. Lamellae under fourth toe 16L/19R (
+holotype
+) or 20/20 (
+paratype
+); terminal three scales above fourth toe undivided.
+
+
+
+Measurements and proportions
+
+
+
+Holotype
+: SVL
+70 mm
+, AGL
+37 mm
+(52.9% of SVL); tail autotomised at base; FLL
+18 mm
+(25.7% of SVL); HLL
+27 mm
+(38.6% of SVL); (FLL + HLL)/AGL 121.6%; HL
+13.4 mm
+(19.1% of SVL); HW 10.0 mm (74.6% of HL); HD
+7.4 mm
+(55.2% of HL).
+
+
+Paratype
+: SVL
+64 mm
+; AGL
+31.5 mm
+(49.2% of SVL); TL
+93 mm
+(145% of SVL); FLL
+18 mm
+(28.1% of SVL); HLL
+27 mm
+(42.2% of SVL); (FLL + HLL)/AGL 142.9%; HL
+13.6 mm
+(21.3% of SVL); HW
+10.1 mm
+(74.3% of HL); HD 8.0 mm (58.8% of HL).
+
+
+Colouration (in preservative).
+Description based on
+holotype
+and
+paratype
+. Dorsal ground colour light olive brown (
+Figs. 3
+&
+4
+). Dorsal head shields with small irregular dark brown spots along some scale margins, particularly medially. Neck and body dorsum with irregular darker brown transversely oriented macules, only weakly aligned to become broken bands, giving appearance of a “chequerboard” pattern of light and dark patches. Dark macules and pale interspaces both about 2–3 scales long from anterior to posterior. Tail dorsum with similar rectangular markings, but less contrasting (mid brown) and more transversely aligned to create bands.
+
+Laterally, ground colour transitioning from light brown dorsally to light yellow-grey ventrally. An irregularly edged dark brown stripe beginning over loreal region (where narrowest), along upper and lower edges of orbit, over upper temporal region (containing a few paler spots), then along upper flanks, where it broadens and breaks up into a coarse dark brown reticulum (containing some spots of pale ground colour) blending with the lateral parts of the dorsal pattern, and extending more ventrally as mid-brown barring. Lateral surface of tail with extensions of the mid-brown dorsal bands.
+
+Venter yellow-grey, throat immaculate, body and tail with mid-brown extensions of the lateral pattern of blotches towards the midline (but not reaching it in the
+paratype
+).
+
+Forelimbs light brown above with weakly defined and coarse brown reticulum above, yellow-grey below; palms yellowish-grey. Hindlimbs light brown above, anteriorly and posteriorly with a coarse dark brown reticulum, yellow-grey below with weak extensions of the dark reticulum encroaching; soles pale yellow.
+Parietal peritoneum black, visceral peritoneum unpigmented.
+
+Colouration (in life).
+Description based on
+holotype
+and unvouchered photographed specimen. Dorsal ground colour light olive brown to golden brown (
+Fig. 5A–C
+). Irregular spotting on dorsal head shields and macules along the dorsum dark brown to black.
+
+Dorsal surface of head sometimes with light greenish tint. Laterally, ground colour transitioning from light brown dorsally to pale ventrally. Venter pale grey. Lining of mouth blue.
+
+
+
+Reproduction.
+Both specimens are female. The
+holotype
+has 1L/2
+R
+thin-shelled eggs in the oviduct indicating viviparity, while the
+paratype
+is non-reproductive but mature (oviduct well-developed with obliquely pleated walls).
+
+
+
+
+Etymology.
+From the Latin masculine noun scurrula, a small clown, in reference to the bright chequerboard pattern, resembling that often used in the dress of clowns and jesters.
+
+
+
+
+Distribution.
+
+Prasinohaema scurrula
+
+
+sp. nov.
+
+is known from three locations spanning approximately
+1,000 km
+2
+of
+Hela Province
+between the Muller Range in the northwest and the Tari area and Gigira Ridge in the southeast (
+Fig. 6
+). The Gigira Ridge (
+-5.972
+,
+142.753
+,
+2,175 m
+a.s.l.) record is based on an unvouchered photograph taken by S. Richards on
+24 June 2015
+(
+Fig. 5A–B
+).
+
+
+
+
+Ecology.
+
+Prasinohaema scurrula
+
+
+sp. nov.
+
+is a montane species. All specimens were found in mid-montane habitats at elevations between
+
+2,175
+–2,900
+m
+
+a.s.l. The
+holotype
+was found on a log in a patch of sun in a fern meadow (
+Fig. 5D–E
+) at Apalu Reke in the Muller Range (see
+Takeuchi 2011
+for a detailed habitat description). The specimen from Gigira Ridge was found on the ground in sunny conditions on an artificially created limestone scree slope at the edge of a road in dense mid-montane forest. Later surveys at the Gigira Ridge location in 2017, 2019, and 2024 did not detect this species. The
+paratype
+was found 40’ (roughly
+12 m
+) high in a pandanus tree (see detailed habitat information in
+Frith & Frith (1990))
+. This suggests arboreal habits for the species, which may explain why observations are rare despite intensive surveys at Apalu Reke and Gigira Ridge, where
+
+Pandanus
+
+trees were common.
+
+
+
+
+
+Prasinohaema scurrula
+.
+
+sp. nov.
+was not observed during a survey at
+500 m
+a.s.l. in the Muller Range, nor at surveys between
+
+1,340
+–2,220
+m
+
+a.s.l. elsewhere in that range (F. Kraus pers. comm.); altitudes below
+2,175 m
+a.s.l. on Gigira Ridge were not sampled.
+
+
+Relatively few other skink species have been observed in sympatry with the new species. An undescribed species of
+
+Papuascincus
+
+(most likely lineage III from
+
+Slavenko
+et al.
+2020
+
+) was abundant at both Apalu Reke and Gigira Ridge. A single unidentified
+
+Emoia
+Gray, 1845
+
+was observed but not captured at Gigira Ridge and a few specimens of an unidentified species of
+
+Lobulia
+
+were also recorded there. This
+
+Lobulia
+
+possibly belongs to an asyet-undescribed species that occurs in the Muller Range, and that is genetically divergent from other species of
+
+Lobulia
+
+present in the Central Cordillera (
+
+Lobulia lobulus
+(
+Loveridge, 1945
+)
+
+and
+
+Lobulia marmorata
+Slavenko, Tamar, Tallowin, Kraus, Allison, Carranza & Meiri, 2021
+
+) (
+
+Slavenko
+et al.
+2022
+
+).
+
+
+
+
+Suggested IUCN conservation status.
+
+Prasinohaema scurrula
+
+
+sp. nov.
+
+is known from three locations spanning approximately
+1,233 km
+2
+of
+Papua
+New Guinea’s Central Cordillera. The Tari Gap and Gigira Ridge locations are respectively about 107 and
+73 km
+east from the Muller Range locality, and separated from each other by ~
+44 km
+. However, these two most proximate localities are separated by the much lower (down to ~
+1,200 m
+a.s.l.) valley of the Tagari River which may represent a significant barrier to dispersal. In contrast, bands of unbroken terrain above
+1,600 m
+a.s.l. connect the Muller Range locality with both Tari Gap and Gigira Ridge. Whether these intervening areas of montane forest and meadow support additional populations of this species is unknown. There are relatively large areas of apparently suitable habitat in the region, and the species has a moderately broad elevational range— suggesting that it is not at imminent risk from climate change. Nonetheless, its distribution, habitat requirements and potential threats remain poorly understood. One striking feature of this species is that at two of the known locations (Apalu Reke and Gigira Ridge), only single individuals were encountered despite intensive surveys lasting at least several days and (at Gigira Ridge) up to a month over several years. In contrast, other montane skinks (
+
+Papuascincus
+spp.
+
+) were abundant at these sites. However, if
+
+P. scurrula
+
+
+sp. nov.
+
+is arboreal, as suggested by the collection details of the
+paratype
+, it would be difficult to detect using terrestrial surveys. Given the moderately broad known range and apparent extensive availability of suitable habitat without major known threats, we recommend that this species should be considered Least Concern.
+
+
+
+
+Comparisons with other species.
+
+Prasinohaema scurrula
+
+
+sp. nov.
+
+differs from all other species of
+
+Prasinohaema
+sensu stricto
+
+in the short blunt snout (vs long, slender and pointed). This is most obvious in lateral profile.Additional features distinguishing it from individual species are covered below.
+
+
+
+Prasinohaema flavipes
+
+is the
+type
+species of the genus and is widely distributed along the Central Cordillera and in the Huon Peninsula.
+
+Prasinohaema scurrula
+
+
+sp. nov.
+
+additionally differs from it in the combination of smaller size (mature individuals
+64–70 mm
+vs
+74.5–104 mm
+), fewer supralabials (seven vs eight or more); lower eyelid with two enlarged opaque scales centrally (vs modally with an opaque to translucent window covering the same area, although some individuals have the window divided into two scales as in
+
+P. scurrula
+
+
+sp. nov.
+
+); basal subdigital lamellae not abruptly broader than distal lamellae (vs basal 7–11 lamellae change abruptly to 7–10 narrow distal lamellae) and with rounded distal edges (vs flat distal edges); temporal scalation without any fragmentation (primary temporal single, upper and lower secondary temporal undivided, last two supralabials undivided) rather than with irregular fragmentation (with at least the primary temporals and last supralabial divided, often with the homologies of the fragmented scales not apparent); parietal scales fully separated by interparietal (vs usually in contact); dorsal colour pattern of alternating dark brown and light brown rectangular regions of roughly equal width in a chequerboard pattern (vs completely banded with dark and light brown regions, or with a broad dorsal stripe or unpatterned;
+Greer & Raizes, 1969
+;
+Kraus, 2010
+); and ventral surface with extensive dark-brown chequerboard pattern (vs plain, or with at most scattered light brown blotching).
+
+
+
+Prasinohaema flavipes paniaiensis
+
+is a subspecies from the Wissel Lakes region of Indonesian New
+Guinea
+that has been largely ignored in the literature since it was described. Geographically isolated from nominate
+
+P. flavipes
+
+in
+Papua New Guinea
+, no evidence was provided at the time of description that it is more closely related to
+
+P. flavipes
+
+than to any of the other
+
+Prasinohaema
+
+, and it has yet to be included in any genetic analyses. It probably warrants recognition at the species level. We provide a comparison based on its original description in
+Brongersma (1949)
+.
+
+Prasinohaema scurrula
+
+
+sp. nov.
+
+shares with
+
+P. flavipes paniaiensis
+
+the separation of the parietal scales by the interparietal, but it differs from it in having the dorsal scales smooth or bluntly grooved (vs with a texture of numerous fine grooves), fewer supralabials (seven vs modally eight), fewer supraciliaries (modally nine vs 9–12, usually 10 or 11), no fragmentation of scales in the temporal region (vs irregular fragmentation), lower eyelid with two large central opaque scales (vs usually a transparent disk, sometimes divided into two or three transparent scales), ear with 2–3 low lobules along anterior margin (vs lacking lobules), basal subdigital lamellae only slightly and gradually diminishing in width distally (vs basally broad, more abruptly transitioning to narrow distal scales), fewer paravertebral scales (65–66 to posterior edge of hindlimbs vs 79–88) and dorsal colour pattern of alternating dark brown and light brown rectangular regions of roughly equal width (vs unbanded, or, if banded, with dark brown and light brown regions of greatly differing width); and ventral surface with extensive dark-brown chequerboard pattern (vs plain, or with at most scattered light brown blotching).
+
+
+
+Prasinohaema prehensicauda
+
+is widely distributed along the Central Cordillera.
+
+P. scurrula
+
+
+sp. nov.
+
+is similar to it in body size (mature-sized individuals
+64–70 mm
+vs
+59–72 mm
+) and in the separation of the parietals by the interparietal.
+
+Prasinohaema scurrula
+
+
+sp. nov.
+
+differs from
+
+P. prehensicauda
+
+in having prefrontals not in contact (vs in contact), fewer supraoculars (4 vs modally 5), fewer supraciliaries (modally nine vs 9–12, usually 10 or more), fewer postsuboculars (4 vs 5–7), fewer supralabials (7 vs 8–10), lower eyelid with two enlarged opaque subpalpebral scales centrally (vs a transparent window formed by a single scale covering the same region), no fragmentation of the temporal region (vs fragmentation), dorsal scales smooth and polished (vs finely shagreened, giving a dull appearance), fewer paravertebrals (65–66 to posterior edge of hindlimbs vs 70–85), longer limbs (FLL/SVL 25.7– 28.1% vs 20.4–25.6%; HLL/SVL 38.6–42.2% vs 25.2–32.3%), more subdigital lamellae (16–20 vs 11–16) and the basal subdigital lamellae only slightly diminishing distally (vs basally broad, more abruptly transitioning to narrow distal scales) and with rounded distal edges (vs flat distal edges).
+
+Prasinohaema prehensicauda
+
+is sexually dichromatic (
+Greer & Raizes 1969
+;
+Woodruff 1972
+;
+Kraus 2010
+). In the absence of male
+
+P. scurrula
+
+
+sp. nov.
+
+, comparison of colouration can only be made with females, but female
+
+P. scurrula
+
+
+sp. nov.
+
+differ from
+
+P. prehensicauda
+
+in having a dorsal pattern of alternating dark brown and light brown rectangular regions of roughly equal anteroposterior width (vs narrow dark bands separated by wider pale intervals containing paler spots).
+
+
+
+Prasinohaema parkeri
+
+is only known from the
+holotype
+, from the Utakwa River in Indonesian New
+Guinea
+, approximately
+500 km
+to the west of the known distribution of
+
+P. scurrula
+
+
+sp. nov.
+
+and has not yet been included in any phylogenetic analysis. Like
+
+P. semoni
+
+and
+
+P. virens
+
+, it may not be part of a monophyletic
+
+Prasinohaema
+
+. However, because it has not yet been excluded from
+
+Prasinohaema
+sensu stricto
+
+, we include a comparison with the new species, based on the description in
+Smith (1937)
+and our examination of the
+holotype
+. The
+holotype
+exhibits a number of unusual scalation features, and it is not known whether these are individual anomalies (part of an apparent pattern in many
+
+Prasinohaema
+species
+
+that exhibit fragmentation or asymmetrical scalation anomalies,
+e.g.
+,
+Brongersma 1949
+) or are character states of the species. We treat these peculiarities as if they characterise the species.
+
+Prasinohaema scurrula
+
+
+sp. nov.
+
+differs from
+
+P. parkeri
+
+in having the anterior loreals distinct, and with partial or complete separation of an upper anterior loreal (vs anterior loreals fused to frontonasals), fewer supralabials (7 vs 8), frontal posteriorly entire, and contacting first two supraoculars, with the frontal overlapping the second supraocular over the entire sutural contact (vs posterior part of frontal partially separated, and this posterior hemiscale contacting (right) or only just separated from (left) the third supraocular, and overlapped by second supraocular); supraoculars unfragmented (vs with fragmentation of the lateral extremities of the second and third supraoculars), supraciliaries 8–9 (vs 13L/12
+R
+), interparietal separating parietals (vs parietals in broad contact posteriorly), temporal region not fragmented (vs fragmented, including division of at least primary temporal and last supralabial), and ventral surface with extensive dark-brown chequerboard pattern (vs plain).
+
+
+
+
\ No newline at end of file
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@@ -0,0 +1,632 @@
+
+
+
+A new species of montane Prasinohaema (Sphenomorphini: Scincidae) from the South-fold Mountains of New Guinea
+
+
+
+Author
+
+Slavenko, Alex
+Cesar Australia, Brunswick, Victoria 3056, Australia
+
+
+
+Author
+
+Shea, Glenn M.
+Sydney School of Veterinary Science, B 01, University of Sydney, New South Wales 2006, Australia & Australian Museum Research Institute, Australian Museum, 10 William St, Sydney, New South Wales 2010, Australia & Herpetology Department, South Australian Museum, North Terrace, Adelaide, South Australia 5000, Australia
+
+
+
+Author
+
+Oliver, Paul M.
+School of Environment and Science, Griffith University, Nathan, Queensland, Australia, and Biodiversity and Geosciences, Queensland Museum, South Brisbane, Queensland 3300, Australia
+
+text
+
+
+Zootaxa
+
+
+2025
+
+2025-04-15
+
+
+5621
+
+
+2
+
+
+211
+230
+
+
+
+
+https://doi.org/10.11646/zootaxa.5621.2.3
+
+journal article
+10.11646/zootaxa.5621.2.3
+1175-5326
+15262507
+B85E0A5A-E44B-4752-B7FF-26202D039517
+
+
+
+
+
+
+Biogeography and evolution of
+
+Prasinohaema
+
+
+
+
+
+
+
+
+Prasinohaema scurrula
+
+
+sp. nov.
+
+is thus far only known from montane areas of the South-fold Mountains in western
+Papua New Guinea
+. While herpetofaunal surveys in these mountains over the last 20 years have resulted in the description of many new montane frog species (e.g.,
+Richards & Oliver 2010
+; Oliver
+et al.
+2012;
+
+Iannella
+et al.
+2014
+
+;
+Oliver & Richards 2019
+;
+Günther & Richards 2021
+),
+
+P. scurrula
+
+
+sp. nov.
+
+is only the fourth new species of montane lizard recognised from this area over the same time period, with the other three being
+
+Cyrtodactylus medioclivus
+Oliver, Richards & Sistrom, 2012
+
+,
+
+Lobulia marmorata
+Slavenko, Tamar, Tallowin, Kraus, Allison, Carranza & Meiri, 2021
+
+and
+
+Papuascincus eldorado
+Slavenko, Richards, Donnellan, Allison & Oliver, 2024
+
+. Of these, three belong to the
+
+Lobulia
+
+group. This contrasting number of frog versus lizard descriptions is likely a cumulative outcome of lower lizard diversity at high elevations (
+
+Tallowin
+et al.
+2017
+
+), a greater research focus on frogs, and potentially more difficulty finding often ecologically cryptic lizard taxa. Two of the three afore-mentioned species (
+
+C. medioclivus
+
+and
+
+P. eldorado
+
+) have wide distributions that span several degrees of longitude but within narrow elevational bands. This pattern of distribution emphasises that for at least some montane taxa there has been connectivity within climatically similar elevational bands spanning large areas of the South-fold Mountains.
+
+
+Lineage diversification and ecological divergence in skinks of the
+
+Lobulia
+
+group appears to be closely associated with mountain uplift and segregation across elevational gradients (
+
+Slavenko
+et al.
+2020
+
+,
+2022
+).
+
+Prasinohaema scurrula
+
+
+sp. nov.
+
+does not appear to be a truly ‘alpine’ species, with no records above 3,000 metres a.s.l. However, it is clearly a montane species and occurs at the highest elevations at which
+
+Prasinohaema
+sensu stricto
+
+has been recorded. The sister genus to
+
+Prasinohaema
+sensu stricto
+
+,
+
+Nubeoscincus
+
+, contains two species, with some of the highest elevational records (~
+
+3,000
+–4,000
+m
+
+a.s.l.) of any skink in New
+Guinea
+, and indeed the world (
+
+Greer
+et al.
+2005
+
+;
+
+Slavenko
+et al.
+2022
+
+). This emphasises the overall close association between the
+
+Lobulia
+
+group, and the
+
+Prasinohaema
+-
+Nubeoscincus
+
+lineage in particular, and mountains. Our divergence dates for
+
+Prasinohaema
+sensu stricto
+
+(~16 Mya) also support models and analyses suggesting that extensive uplift in at least some parts of what is now New Guinea’s Central Cordillera date back well into the Miocene (
+Quarles van Ufford & Cloos 2005
+;
+
+Tallowin
+et al.
+2018
+
+;
+
+Slavenko
+et al.
+2020
+
+,
+2022
+;
+
+Toussaint
+et al.
+2021
+
+).
+
+
+
+
+When compared against its congeners
+
+P. scurrula
+
+
+sp. nov.
+
+has a relatively robust build and less extensive lamellae on the digits, suggesting more terrestrial habits. However, the
+paratype
+was collected high up in a
+
+Pandanus
+
+tree, suggesting arboreality. The two other observations we have of this species in the wild were of lizards on or close to the ground, but at forest edges or clearings. The morphology of
+
+P. scurrula
+
+
+sp. nov.
+
+is therefore reminiscent of arboreal lizards such as the ‘twig anole’ ecomorph, which is characterised by short limbs and tails and small lamellae (
+Williams 1983
+;
+Losos 2009
+), rather than trunk dwelling species.
+
+
+In New Guinea’s higher mountains (>
+2,500 m
+a.s.l.) largely terrestrial skink lineages such as
+
+Carlia
+Gray, 1845
+spp.
+
+,
+
+Emoia
+spp.
+
+and other radiations of sphenomorphin skinks currently assigned to the genus
+
+Sphenomorphus
+Fitzinger, 1843
+
+are relatively rare or absent. In contrast arboreal and semi-arboreal species such as
+
+Lobulia
+spp.
+
+,
+
+Prasinohaema
+sensu stricto
+
+spp. (including the new species described here), and some members of
+
+Ornithuroscincus
+
+dominate. The elevational distributions of these different skink radiations may provide evidence to support the argument that vertebrate lineages in the tropics will often show increasing arboreality as elevation increases (
+
+Scheffers
+et al.
+2013
+
+). The basis of this argument is that there is generally little overlap in temperature regimes between low- and high-elevation ground and soil microhabitats (
+Scheffers & Williams 2018
+), meaning that lowland terrestrial species are unlikely to be able to extend to high elevations in the tropics (
+
+Scheffers
+et al.
+2013
+
+). In contrast, arboreal tropical species are likely to have broader physiological tolerances and wider distributions (
+Withers 1981
+;
+Adolph 1990
+;
+Wikramanayake & Dryden 1993
+). Based on these observations,
+Scheffers & Williams (2018)
+concluded that arboreal lineages are more likely to have broader niches, higher dispersal capabilities (
+
+Scheffers
+et al.
+2017
+
+) and increased diversification rates (
+Moen & Wiens 2017
+). Greater physiological tolerance in the more arboreal
+
+Lobulia
+
+group is one potential explanation for the greater success of this lineage in montane habitat. In turn, ecological release could have allowed members of the more arboreal
+
+Lobulia
+
+group to diversify and exploit vacant terrestrial niches once they colonised montane areas. We suspect this may be the case both for the true ‘alpine’ species—such as
+
+Alpinoscincus
+spp.
+
+and
+
+Nubeoscincus
+spp.
+
+, which occupy rocky outcrops, clearings and grasslands above the tree line (
+
+Greer
+et al.
+2005
+
+)—and in the heliothermic and basking
+
+Papuascincus
+spp.
+
+To test this hypothesis further a more formal analysis of the evolutionary trajectory, historical biogeography and correlates of major ecological shifts in the
+
+Lobulia
+
+group would be needed, combined with more rigorous studies on the ecophysiology of New Guinea’s montane skinks. By better documenting the species diversity of this radiation we are hopefully improving the framework needed for such an analysis.
+
+
+The description of
+
+P. scurrula
+
+
+sp. nov.
+
+also emphasises the many residual issues associated with the taxonomy of
+
+Prasinohaema
+
+.
+Greer (1974)
+originally defined the genus largely based on the presence of a prehensile tail with a glandular tip and green blood plasma due to extremely high concentrations of biliverdin (
+Greer & Raizes 1969
+;
+Austin & Jessing 1994
+). However, Greer recognised that the genus shows variation in reproductive mode and the condition of the secondary palate. With the advent of molecular phylogenetics, it has become evident that the genus
+
+Prasinohaema
+
+is not monophyletic and that key traits—including green blood plasma—probably evolved several times independently (
+
+Rodriguez
+et al.
+2018
+
+;
+
+Slavenko
+et al.
+2022
+
+), meaning a revised diagnosis of the major lineages of so-called “green-blooded” skinks is needed. Additionally, Rodriquez
+et al.
+(2018) provided evidence of unrecognised taxa in both
+
+Prasinohaema
+sensu stricto
+
+and
+
+Prasinohaema
+sensu lato
+virens
+
+. It has also been known for decades that
+
+Prasinohaema
+sensu lato
+semoni
+
+likely comprises a species complex (G. Shea pers. obs.), and we are aware of additional morphologically distinctive populations of named species that have as yet not been subject to genetic analysis. As noted above the status of
+
+P. flavipes paniaiensis
+
+remains unresolved, but it seems likely that this form is not conspecific with
+
+P. flavipes
+
+; indeed it shares several similarities with
+
+P. prehensicauda
+
+. Our genetic data build on this background by further emphasising deep genetic divergences between populations ascribed to the nominal taxa
+
+P. flavipes
+
+and
+
+P. prehensicauda
+
+and highlighting a need for detailed taxonomic investigations of these taxa. This in turn highlights the broader pattern that skinks are both the most diverse, and the most poorly known, of Melanesia’s major reptile radiations (
+
+Slavenko
+et al.
+2023
+
+). Addressing these knowledge gaps is important to improve our understanding of Melanesian biogeographic history, and to better elucidate centres of diversity and endemism across the region.
+
+
+
+Nomenclature of skink scales
+
+
+
+The large variation in definitions of skink scales, particularly on the head and around the eyes, has created issues when comparing states across different publications. We provide here a thorough overview of different scale definitions used in skinks, particularly in the Australasian region, and stabilise the nomenclature. An accompanying labelled diagram of the scale definitions we used can be found in
+Figure 2
+.
+
+
+Taylor (1936)
+defines the supraciliaries as being the line of small scales bordering the lateral margin of the supraocular scales, from the first scale contacting the prefrontal to the last scale wedging between the last supraocular and parietal, and overlapping the uppermost postsubocular, and this definition was followed by us and, among others,
+Shea & Greer (2002)
+,
+Zug (2004)
+,
+Das & Austin (2007)
+,
+Kraus (2007
+, 2018, 2020),
+Shea (2017)
+and
+Shea & Allison (2021)
+in the Indopapuan region. Others (
+e.g.
+,
+Hardy 1977
+;
+Horner 2007
+;
+
+Grismer
+et al.
+2011
+
+;
+Hedges & Conn 2012
+) exclude the posteriormost supraciliary, either leaving it unnamed (
+Horner 2007
+), or referring to it as an anterior postocular (
+Hardy 1977
+), an upper secondary postocular (
+Hedges & Conn 2012
+), an anterior primary temporal (
+
+Grismer
+et al.
+2011
+
+) or an anterior pretemporal (
+Greer 1983
+;
+
+Greer
+et al.
+2005
+
+; followed by
+
+Slavenko
+et al.
+2022
+
+,
+2024
+).
+
+
+Greer’s (1983)
+creation of the name pretemporals for the posteriormost supraciliary and posteriormost postsubocular of Taylor’s nomenclature was originally due to the combined fate of these two scales in some Australian
+
+Lerista
+Bell, 1833
+species
+
+, in which they are fused into a single scale. These scales are not fused in New
+Guinea
+skinks, making the term pretemporals superfluous in this geographical context.
+
+
+Other definitional variation exists at the anterior end of the row of supraciliaries.
+Hardy (1977)
+, working on
+New Zealand
+skinks, excluded the first supraciliary from the count, identifying this as an upper preocular.
+
+
+Taylor (1936)
+identified as presuboculars the row of scales bordering the dorsal margins of the supralabial scales, from the first scale posterior to the second loreal, to the last scale being that which reaches the supralabial that lies under the centre of the eye (the subocular supralabial), and the preocular being the scale anterior to both the upper and lower palpebral rows that lies dorsal to the first presubocular, ventral to the supraciliaries, and posterior to the second loreal, sometimes succeeded by a smaller scale posteriorly (
+e.g.
+, Greer 1982). Others (
+e.g.
+,
+Greer & Shea 2004
+;
+
+Greer
+et al.
+2005
+
+;
+
+Slavenko
+et al.
+2022
+
+,
+2024
+working with New
+Guinea
+skinks) consider the first presubocular of Taylor’s definition to be the lower of two preoculars. To convert from Taylor’s definition to that used by
+Greer & Shea (2004)
+, the number of preoculars needs to be increased by one, and the number of presuboculars decreased by one.
+Taylor (1936)
+defines the postsuboculars as the line of scales bordering the posteroventral margin of the orbit (and overlying the anterior edge of the jugal bone), from the first scale contacting both the subocular supralabial and the supralabial behind that, up to and including the scale wedging behind the last supraciliary and parietal, while postoculars are the slightly enlarged scales immediately behind the upper and lower palpebral margins.
+Greer (1983)
+reidentified the last of Taylor’s postsuboculars as the second of his pretemporals, while
+Greer & Shea (2004)
+considered only those of Taylor’s postsuboculars that contacted the penultimate supralabial as being true postsuboculars, thereby excluding some of Taylor’s scales, which Greer does not otherwise discuss in his descriptions. However, as many of the New Guinean sphenomorphins have divisions of the penultimate supralabial, using this scale to define the postsuboculars risks counts on species with varying degrees of division of the supralabial not being homologous. Here we favoured Taylor’s postsubocular definition since it is consistent with the underlying jugal bone.
+
+
+Paravertebral scales also have a wide range of definitions in the literature. While most begin their count from the first scale behind the parietals, some (
+e.g.
+,
+
+Amarasinghe
+et al.
+2016
+
+;
+
+Slavenko
+et al.
+2022
+
+) exclude the first nuchal scale (the primary nuchal of
+Miralles 2006
+) from the count, and others (
+e.g.
+,
+Feria-Ortiz & Garcia-Vázquez 2012
+) exclude all nuchals from the count. The posterior extremity of the count is even less consistently defined across studies. Some authors count to the last scale anterior to the hindlimb (
+e.g.
+,
+Shea & Greer 2002
+;
+Shea & Allison 2021
+;
+
+Slavenko
+et al.
+2022
+
+) or crossing a line at the level of the anterior margin of the hindlimbs (
+
+Slavenko
+et al.
+2024
+
+), others count back to the last scale anterior to the posterior margin of the hindlimb (
+e.g.
+,
+Zug 2004
+), the last scale at the midpoint between the posterior margins of the hindlimbs (
+e.g.
+,
+Horner 2007
+), the last scale immediately posterior to the posterior margin of the hindlimb (
+e.g.
+,
+
+Greer
+et al.
+2005
+
+), the last scale opposite the vent (
+e.g.
+,
+
+Grismer
+et al.
+2011
+
+;
+Hedges & Conn 2012
+) or “the base of the tail”, without defining that point (
+e.g.
+,
+Das & Austin 2007
+). We provided two counts to cover some of this variation, which should allow future taxonomists easier comparison to at least one of our counts.
+
+
+
+
\ No newline at end of file