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Mokopirirakau galaxias sp. nov. (Figs 2���5) Holotype. Adult male NZMZ RE.008613, from an undisclosed location within Oteake Conservation Park, North Otago, New Zealand, collected by C. Knox and T. Jewell on 14 January 2020. The collection site (and all other known populations of Mokopirirakau galaxias sp. nov.) are between latitudes of 44�� 35���S���44�� 58���S, longitudes of 169�� 43���E���170�� 19���E, and altitudes of 1,300-1,600 m ASL. Paratypes. None (but see Variation section below). Etymology. Latin (from Ancient Greek) galaxias for the patterning of small white spots on the dorsal and lateral surfaces of the body reminiscent of the Milky Way. The epithet is formed as a noun in apposition. Suggested Common Name. ���Hura te ao gecko.��� From te reo M��ori, the indigenous language of New Zealand���s first people, for ���breaking dawn.��� ���Hura��� commonly means ���reveal,��� but can also be applied colloquially to signify ���the break of dawn.��� This concept ties in with aspects of this species��� colouration (i.e., the orange-yellow around the mouth, evocative of sunlight at daybreak), as well as the idea that this lizard has for the first time revealed itself to us. Diagnosis. Both the phylogenetic analysis and the combination of straight rather than strongly curved or vshaped lamellae, presence of brightly coloured mouth pigment, absence of apical plates from some toes, lamellae forming a pad which does not extend to the end of the toe, and ventral scales on the tail being roughly twice the size of dorsal scales, identify the species as a member of the genus Mokopirirakau. The species can readily be distinguished from all Mokopirirakau species, except M. kahutarae, on the basis of eye colour (very dark brown versus grey, light brown or occasionally light blue with dark ���filigree��� markings), body colour pattern (olive-grey to olive-green with small white spots versus grey with transverse bands), and head colour pattern (contrasting v-shaped dark marking behind eyes and pale stripe from eye to ear absent in this species and M. kahutarae; present in all others). Distinguished from M. kahutarae, to which it has some obvious similarities (see Whitaker 1984), on the basis of a number of characters: on average smaller body size (76���84 mm SVL versus up to 91 mm); snout slightly more elongate (longer than versus equal to eye-ear distance); tail length shorter than SVL (versus equal or longer); distal attenuation of digits ~25% (versus 31%), with webbing extending to approximately half of the distal phalange (versus approximately one quarter); 7���9 subdigital lamellae (versus 12���14); apical plates slightly enlarged on the first 3 digits of each autopodium (versus absent); single scale row between middle of orbit and labial scales (versus 2 rows); 2���3 supraciliary scale rows (versus 4���5), forming an ���eyebrow,��� with supraciliary scales scarcely enlarged (versus greatly enlarged), with the effect of upper margin of eye fully visible from the side (versus hidden by an ���eyebrow���); 8���9 infralabial scales (versus 11���12); ~18 ventral scale rows from axilla to groin (versus ~24); two (versus three) cloacal spurs; iris very dark brown (versus totally black); pupil margin very dark brown (versus black); labial scales and above surrounding scales, as well as the palmar surfaces of the manus, pes, and digits coloured vivid orange (versus muted olive-grey); mouth and throat colour orange-pink and grey-black, respectively (versus pink to pale red); base colouration of dorsal skin more consistently olive (versus olive to slate- grey); and dorsal markings composed of numerous pale spots with narrow, asymmetrical paler transverse dorsal markings (sometimes absent or indistinct) (versus 6���7 wide pale grey bands). Table 1 compares characters of M. galaxias sp. nov. with all other described Mokopirirakau species. Diagnostics are not given for any other non-described Mokopirirakau species on the basis that formal taxonomic descriptions for these are in preparation (R. Hitchmough, S.V. Nielsen and A. Bauer, unpub. data); however, none of these resemble M. galaxias and all are allopatrically distributed. Description of holotype. Adult male, 68 mm SVL, 65 mm TL. Dorsal scales fine, regular. Ventral scales regular, larger than dorsal. Tail cylindrical, tapering. Eye large. Snout blunt. Rostral broader than long, with deep cleft, c. 50% of scale. Nasals 3. Internasals 1, Supralabials/upper labials 11. Infralabials/lower labials 9. Mental 1, broader than wide. Postmentals 1. Precloacal pores. 20 wide, 4 deep. Cloacal spurs 2. Digits moderate, narrow. Lamellae 9, straight. Measurements (in mm): HL 23.5; HW 15.5; AG 31; SF 29.5; S-E 18; ED 3.8; E-E 10.2; EF 12.5. Ratios: HL/ED 6.18. SE/EE 1.76; TL/SVL 0.95. Colour. Base body colour olive-grey to olive-green on the dorsum with a light grey venter. Dorsal markings indistinct, consisting of thin blotches and/or streaks extending outward from the midline or slightly alternating and overlapping the midline, markings coalesce to form a broader patch on top of head, and continue down tail. Small, whitish spots on dorsal and particularly numerous along the flanks, sometimes aligning to form secondary, transverse bars between main dorsal markings; spots vary in intensity between individuals, but often dominate the overall colour pattern. Head markings subtle, indiscernible or absent. Eye particularly large, lined with orange, almost touching labial scales, with enlarged supraciliary fringe creating slightly ���hooded��� eyebrow. Iris dark blackish brown; black pupil with paler edges just discernible. Labials and adjacent scales distinctively pale yellow-olive to orange. Tongue light pink in middle, darker stippled pink at tip, with bright orange base. Roof of mouth bright orange on and around gums; coloured from pink anteriorly to mid-grey to black at back and in throat. Floor of mouth pink, with bright orange at base of tongue and thin lining along inner margin of gums. Ventral very finely peppered with lighter and darker specks. Cloacal opening bright orange. Soles and undersurface of digits pale to bold creamyorange. Variation. Twenty one live individuals (eight adult females, five adult males, four subadult females, three subadult males, and one juvenile) were captured, measured, photographed, and released across the four known sites between January 2018 and December 2020. The following data is from the first six specimens captured: adult males, adult females; 76���84 mm SVL (x=80, SD 3.5). Dorsal and ventral scales as for holotype. Tail cylindrical, tapering. Eye large. Snout blunt. Rostral, broader than long, with deep cleft c. 50% of scale. Nasals 3 or 4. Internasals 0���2. Supralabials/upper labials 9���11, Infralabials/lower labials 8���9, usually 9. Mental 1. Postmentals 1. Precloacal pores 21���24 wide, 4���5 deep. Cloacal spurs 2���4 in males only. Digits moderate, narrow. Lamellae 7���9. straight. Ratios: HL/ED 4.2���4.5 (0.15), SE/EE1.10���1.28 (0.07), TL/SVL 0.83���0.89 (0.02). Twenty specimens had intact original tails and these were consistently much shorter than the snout-to-vent length. The exception was an adult male for which almost the whole tail was regenerated, yet was still the same proportional length as for those with their original tails. Distribution. Mokopirirakau galaxias sp. nov. appears to be an alpine specialist and is known only from Oteake Conservation Park. This species is currently known from c.8 rock tors and bluffs within the 1.4-ha type locality, plus three additional sites in deep boulderfield habitat. It could possibly occur elsewhere in Otago and Canterbury, despite different rock types and other Mokopirirakau species being present in some areas, if sufficient niche separation is possible. Both M. kahutarae and M. galaxias are thought to be solely alpine in nature; however, distributional limits and pre-human altitudinal limits of both are unknown. The ~400-km distributional gap between them may at least partly be an artefact of poor sampling, particularly given the difficult access to many of their habitats. Suitably creviced rock exposures, deep boulderfields and talus slopes occur widely in the mountain ranges between their known distributions but are patchy in distribution. Environment. The climate in the 64.03 Hawkdun and 64.04 Saint Bathans Ecological Districts is semi-arid with warm, dry summers and cold winters (McEwen 1987). LENZ Level IV data indicate the land environment at the type locality is particularly harsh, with an annual average temperature of 4.2 ��C, a mean minimum temperature of -4.1 ��C during the coldest month of the austral winter (July), average annual solar radiation of 13.2 MJ/m2/day and winter solar radiation of 3.8 MJ/m2 /day (Leathwick et al. 2002). Habitat. Fractured and deeply creviced greywacke outcrops and deep boulderfields and talus slopes. Snow tussock (Chionochloa rigida) grassland and Dracophyllum rosmarinifolium shrubland are the predominant vegetation types in the areas where the species has been found, interrupted by greywacke rock outcrops or boulderfields and talus. At the type locality, M. galaxias have been found on six discrete, but proximate rock outcrops and bluffs comprised of a protruding but deeply fissured and fractured central tor several metres tall, surrounded by varying amounts of rock talus, fellfield and/or scree, along a ~270-m section of a ridgeline. The first gecko discovered was located within a crevice inside a c. 50 cm fractured rock, in an expanse of stony ground c. 10 m from a major outcrop system. The other eighteen geckos encountered at the type locality have all been inside vertical rock fissures, or out on rock ledges. At the other three sites (across two mountain ranges) the five geckos observed were emergent in deep boulderfield or talus, but three of these were well concealed by surrounding rocks and were in positions close to retreats or escape routes. Some of the rocks at all known sites, particularly on tors, are heavily encrusted with colourful lichens of various species, including large amounts of the tall, bristly Neuropogon spp., the latter which may limit the extent of usable rock surface for the geckos (Fig. 6). Natural history. Mokopirirakau galaxias is apparently an alpine specialist gecko, saxicolous in greywacke rock outcrops, boulderfields, and talus. The geckos are strongly nocturnal and thigmothermic. They have been detected on exposed rock surfaces during surveys at night in air temperatures of 9-16 ��C. Nothing is known of the species��� life history, diet or reproduction. The sex ratio appears to be close to 1:1 based on the limited survey results to date (C. Knox, unpub. data). A small number of juvenile and subadult geckos have been located, but not neonates. It is likely to be mostly insectivorous, but may consume berries of alpine shrubs where available. It is likely to be viviparous, as all New Zealand geckos are. However, due to the species��� alpine niche, it is most likely slow-growing, late to mature (at least 5 years) and long lived (to decadal timeframes), with a very low reproductive output (Cree 1994, Cree & Hare 2016; Knox et al. 2019). One of two adult females captured in January 2018 and a female captured in November 2020 were pregnant; however, no inference can be made on the annual reproductive output per female based on this very small sample size. At all known sites M. galaxias is sympatric with the Southern Alps gecko (Woodworthia ���Southern Alps���), which is sometimes numerous and occupies all available rocky habitat from outcrops to talus and scree. The two gecko species can be syntopic, sharing the same rock crevice or foraging on the same rock ledge. Size differences between M. galaxias (to 84 mm SVL) and W. ���Southern Alps��� (c. 55-65 mm SVL) may allow resource partitioning in retreat site and prey selection between the two species. In contrast, Whitaker et al. (1999) noted that in Nelson and Marlborough, M. kahutarae appear to exclude specimens of the sympatric Woodworthia geckos from their immediate territory. Several species of Oligosoma skinks are also present, particularly O. maccanni, which are found throughout the same rocky habitat and in the surrounding alpine tussockland. Phylogenetic affinities. Genetic analyses using sequences from the mitochondrial ND2 gene indicate that M. galaxias is sister to a clade comprising the sister species M. granulatus and M. kahutarae. Mokopirirakau granulatus occurs in the upper half of the North Island, and in Marlborough, Tasman, and Westland in the South Island. The closest M. kahutarae populations to M. galaxias are approximately 400 km to the north-east; whereas, the closest M. granulatus populations to M. galaxias are approximately 180 km away, but on the other side of the Southern Alps (Fig. 7). Thus, M. galaxias is completely isolated from its nearest relatives geographically, with multiple mountain ranges between their known ranges. The geographically closest Mokopirirakau populations are of the undescribed taxon M. ���Roy���s Peak���, which occurs to the west and south-west (van Winkel et al. 2018; Knox et al. 2019). Conservation. At the type locality this species is currently known only from eight rock tors and surrounding talus within a 1.4-ha area, and 10 ��C, dry and calm nights), but did not locate new populations. However, in November and December 2020, three new populations were found, including two sites in another mountain range within Oteake Conservation Park. Given the November and December 2020 finds, it is deemed likely that other populations of this species will be discovered, as further survey work is undertaken in North Otago and South Canterbury. Threats to M. galaxias include predation from introduced mammals, illegal collection, and climate change. In addition, fires and grazing by introduced herbivores may be a threat by reducing habitat quality and food availability. Introduced mammalian carnivores and rodents that may encroach into their high altitude, exposed habitat pose a threat, especially predators that hunt at night when the geckos may be emergent on rock surfaces. These are likely to include cats (Felix catus), ferrets (Mustela furo), stoats (Mustela erminea), weasels (Mustela nivalis), hedgehogs (Erinaceus europaeus), possums (Trichosurus vulpecula), and mice (Mus musculus) (O���Donnell et al. 2017; Josh Kemp pers. comm.). Mice are likely to have population irruptions following tussock mast seeding, which will then drive secondary carnivore irruptions. Stoats are known to prey on alpine lizards (O���Donnell et al. 2017), and mice and weasels can squeeze into tight crevices to access the geckos inside retreats. Illegal collection has emerged as a threat to several species of New Zealand lizards, particularly geckos (Knox et al. 2013; Marshall et al. 2020) and if precise site information becomes publicly available then determined smugglers could impact the abundance of geckos at those sites. Climate change may also render any remaining habitat increasingly vulnerable to introduced mammals, as it is expected that more predators will infiltrate alpine areas as the climate warms (O���Donnell et al. 2017). Due to the species being discovered very recently, it is not yet listed in the most recent iteration of the Department of Conservation���s New Zealand Threat Classification System lists (Hitchmough et al. 2016). Typically, newly discovered species are placed in the ���Data Deficient��� category. However, the system requires a precautionary ap- proach under which, provided some information exists, the species is placed into the most threatened category consistent with what is known, with uncertainty being indicated by a ���Data Poor��� qualifier. We recommend the species be assigned the conservation ranking of ���Threatened���Nationally Endangered��� with the qualifier ���Data Poor��� (Townsend et al. 2008). This is because of its low abundance, likely decline resulting from impacts of predatory mammals, threats from climate change, and highly restricted range (four small locations). This species should be a high priority for the Department of Conservation for surveys to improve understanding of its distribution, conservation status and conservation management requirements. In the IUCN Red List system, in the absence of clear information on population trend, Data Deficient seems the best recommendation for listing., Published as part of Knox, Carey, Hitchmough, Rodney A., Nielsen, Stuart V., Jewell, Tony & Bell, Trent, 2021, A new, enigmatic species of black-eyed gecko (Reptilia: Diplodactylidae Mokopirirakau) from North Otago, New Zealand, pp. 140-156 in Zootaxa 4964 (1) on pages 145-153, DOI: 10.11646/zootaxa.4964.1.7, http://zenodo.org/record/4706799, {"references":["Whitaker, A. H. (1984) Hoplodactylus kahutarae n. sp. (Reptilia; Gekkonidae) from the Seaward Kaikoura Range, Marlborough, New Zealand. New Zealand Journal of Zoology, 11, 259 - 270. https: // doi. org / 10.1080 / 03014223.1984.10428239","McEwen, W. M. (1987) Ecological Regions and Districts of New Zealand (4 volumes). New Zealand Biological Resources Centre, Department of Conservation.","Leathwick, J., Morgan, F., Wilson, G., Rutledge, D., McLeod, M. & Johnston, K. (2002) Land Environments of New Zealand: A Technical Guide. Ministry for the Environment, Hamilton, 244 pp.","Cree, A. (1994) Low annual reproductive output in female reptiles from New Zealand. New Zealand Journal of Zoology, 21 (4), 351 - 372. https: // doi. org / 10.1080 / 03014223.1994.9518005","Cree, A. & Hare, K. M. (2016) Reproduction and life history of New Zealand lizards. In: Chapple, D. G. (Ed.), New Zealand Lizards. Springer, Berlin, pp. 169 - 206. https: // doi. org / 10.1007 / 978 - 3 - 319 - 41674 - 8 _ 7","Knox, C. D., Jewell, T. R. & Monks, J. M. (2019) Ecology of orange-spotted geckos (Mokopirirakau \" Roys Peak \") in Central Otago and Queenstown-Lakes districts. New Zealand Journal of Ecology, 43 (2), 1 - 9. https: // doi. org / 10.20417 / nzjecol. 43.14","Whitaker, T [A. H.]., Shaw, T. & Hitchmough, R. (1999) Black-eyed geckos (Hoplodactylus kahutarae) on Mt Arthur, Kahurangi National Park. Conservation Advisory Science Notes No. 230. [Department of Conservation, Wellington.]","O'Donnell, C. F. G., Weston, K. A. & Monks, J. M. (2017) Impacts of introduced mammalian predators on New Zealand's alpine fauna. New Zealand Journal of Ecology, 41 (1), 1 - 22. https: // doi. org / 10.20417 / nzjecol. 41.18","Knox, C. D., Cree, A. & Seddon, P. J. (2013) Accurate identification of individual geckos (Naultinus gemmeus) through dorsal pattern differentiation. New Zealand Journal of Ecology, 37, 60 - 66.","Marshall, B. M., Strine, C. & Hughes, A. C. (2020) Thousands of reptile species threatened by under-regulated global trade. Nature Communications, 11 (1), 1 - 12. https: // doi. org / 10.1038 / s 41467 - 020 - 18523 - 4","Hitchmough, R., Barr, B., Monks, J., Lettink, M., Reardon, J., Tocher, M., van Winkel, D. & Rolfe, J. (2016) Conservation status of New Zealand reptiles, 2015. New Zealand Threat Classification Series [Department of Conservation, Wellington, New Zealand], (17), 14 pp.","Townsend, A. J., de Lange, P. J., Duffy, C. A. J, Miskelly, C. M., Molloy, J. & Norton, D. A. (2008) New Zealand Threat Classification System Manual. Department of Conservation, Wellington, 35 pp."]}

Mokopirirakau galaxias sp. nov. (Figs 2–5) Holotype. Adult male NZMZ RE.008613, from an undisclosed location within Oteake Conservation Park, North Otago, New Zealand, collected by C. Knox and T. Jewell on 14 January 2020. The collection site (and all other known populations of Mokopirirakau galaxias sp. nov.) are between latitudes of 44º 35’S–44º 58’S, longitudes of 169º 43’E–170º 19’E, and altitudes of 1,300-1,600 m ASL. Paratypes. None (but see Variation section below). Etymology. Latin (from Ancient Greek) galaxias for the patterning of small white spots on the dorsal and lateral surfaces of the body reminiscent of the Milky Way. The epithet is formed as a noun in apposition. Suggested Common Name. ‘Hura te ao gecko.’ From te reo Mâori, the indigenous language of New Zealand’s first people, for ‘breaking dawn.’ ‘Hura’ commonly means ‘reveal,’ but can also be applied colloquially to signify ‘the break of dawn.’ This concept ties in with aspects of this species’ colouration (i.e., the orange-yellow around the mouth, evocative of sunlight at daybreak), as well as the idea that this lizard has for the first time revealed itself to us. Diagnosis. Both the phylogenetic analysis and the combination of straight rather than strongly curved or vshaped lamellae, presence of brightly coloured mouth pigment, absence of apical plates from some toes, lamellae forming a pad which does not extend to the end of the toe, and ventral scales on the tail being roughly twice the size of dorsal scales, identify the species as a member of the genus Mokopirirakau. The species can readily be distinguished from all Mokopirirakau species, except M. kahutarae, on the basis of eye colour (very dark brown versus grey, light brown or occasionally light blue with dark ‘filigree” markings), body colour pattern (olive-grey to olive-green with small white spots versus grey with transverse bands), and head colour pattern (contrasting v-shaped dark marking behind eyes and pale stripe from eye to ear absent in this species and M. kahutarae; present in all others). Distinguished from M. kahutarae, to which it has some obvious similarities (see Whitaker 1984), on the basis of a number of characters: on average smaller body size (76–84 mm SVL versus up to 91 mm); snout slightly more elongate (longer than versus equal to eye-ear distance); tail length shorter than SVL (versus equal or longer); distal attenuation of digits ~25% (versus 31%), with webbing extending to approximately half of the distal phalange (versus approximately one quarter); 7–9 subdigital lamellae (versus 12–14); apical plates slightly enlarged on the first 3 digits of each autopodium (versus absent); single scale row between middle of orbit and labial scales (versus 2 rows); 2–3 supraciliary scale rows (versus 4–5), forming an ‘eyebrow,’ with supraciliary scales scarcely enlarged (versus greatly enlarged), with the effect of upper margin of eye fully visible from the side (versus hidden by an ‘eyebrow’); 8–9 infralabial scales (versus 11–12); ~18 ventral scale rows from axilla to groin (versus ~24); two (versus three) cloacal spurs; iris very dark brown (versus totally black); pupil margin very dark brown (versus black); labial scales and above surrounding scales, as well as the palmar surfaces of the manus, pes, and digits coloured vivid orange (versus muted olive-grey); mouth and throat colour orange-pink and grey-black, respectively (versus pink to pale red); base colouration of dorsal skin more consistently olive (versus olive to slate- grey); and dorsal markings composed of numerous pale spots with narrow, asymmetrical paler transverse dorsal markings (sometimes absent or indistinct) (versus 6–7 wide pale grey bands). Table 1 compares characters of M. galaxias sp. nov. with all other described Mokopirirakau species. Diagnostics are not given for any other non-described Mokopirirakau species on the basis that formal taxonomic descriptions for these are in preparation (R. Hitchmough, S.V. Nielsen and A. Bauer, unpub. data); however, none of these resemble M. galaxias and all are allopatrically distributed. Description of holotype. Adult male, 68 mm SVL, 65 mm TL. Dorsal scales fine, regular. Ventral scales regular, larger than dorsal. Tail cylindrical, tapering. Eye large. Snout blunt. Rostral broader than long, with deep cleft, c. 50% of scale. Nasals 3. Internasals 1, Supralabials/upper labials 11. Infralabials/lower labials 9. Mental 1, broader than wide. Postmentals 1. Precloacal pores. 20 wide, 4 deep. Cloacal spurs 2. Digits moderate, narrow. Lamellae 9, straight. Measurements (in mm): HL 23.5; HW 15.5; AG 31; SF 29.5; S-E 18; ED 3.8; E-E 10.2; EF 12.5. Ratios: HL/ED 6.18. SE/EE 1.76; TL/SVL 0.95. Colour. Base body colour olive-grey to olive-green on the dorsum with a light grey venter. Dorsal markings indistinct, consisting of thin blotches and/or streaks extending outward from the midline or slightly alternating and overlapping the midline, markings coalesce to form a broader patch on top of head, and continue down tail. Small, whitish spots on dorsal and particularly numerous along the flanks, sometimes aligning to form secondary, transverse bars between main dorsal markings; spots vary in intensity between individuals, but often dominate the overall colour pattern. Head markings subtle, indiscernible or absent. Eye particularly large, lined with orange, almost touching labial scales, with enlarged supraciliary fringe creating slightly ‘hooded’ eyebrow. Iris dark blackish brown; black pupil with paler edges just discernible. Labials and adjacent scales distinctively pale yellow-olive to orange. Tongue light pink in middle, darker stippled pink at tip, with bright orange base. Roof of mouth bright orange on and around gums; coloured from pink anteriorly to mid-grey to black at back and in throat. Floor of mouth pink, with bright orange at base of tongue and thin lining along inner margin of gums. Ventral very finely peppered with lighter and darker specks. Cloacal opening bright orange. Soles and undersurface of digits pale to bold creamyorange. Variation. Twenty one live individuals (eight adult females, five adult males, four subadult females, three subadult males, and one juvenile) were captured, measured, photographed, and released across the four known sites between January 2018 and December 2020. The following data is from the first six specimens captured: adult males, adult females; 76–84 mm SVL (x=80, SD 3.5). Dorsal and ventral scales as for holotype. Tail cylindrical, tapering. Eye large. Snout blunt. Rostral, broader than long, with deep cleft c. 50% of scale. Nasals 3 or 4. Internasals 0–2. Supralabials/upper labials 9–11, Infralabials/lower labials 8–9, usually 9. Mental 1. Postmentals 1. Precloacal pores 21–24 wide, 4–5 deep. Cloacal spurs 2–4 in males only. Digits moderate, narrow. Lamellae 7–9. straight. Ratios: HL/ED 4.2–4.5 (0.15), SE/EE1.10–1.28 (0.07), TL/SVL 0.83–0.89 (0.02). Twenty specimens had intact original tails and these were consistently much shorter than the snout-to-vent length. The exception was an adult male for which almost the whole tail was regenerated, yet was still the same proportional length as for those with their original tails. Distribution. Mokopirirakau galaxias sp. nov. appears to be an alpine specialist and is known only from Oteake Conservation Park. This species is currently known from c.8 rock tors and bluffs within the 1.4-ha type locality, plus three additional sites in deep boulderfield habitat. It could possibly occur elsewhere in Otago and Canterbury, despite different rock types and other Mokopirirakau species being present in some areas, if sufficient niche separation is possible. Both M. kahutarae and M. galaxias are thought to be solely alpine in nature; however, distributional limits and pre-human altitudinal limits of both are unknown. The ~400-km distributional gap between them may at least partly be an artefact of poor sampling, particularly given the difficult access to many of their habitats. Suitably creviced rock exposures, deep boulderfields and talus slopes occur widely in the mountain ranges between their known distributions but are patchy in distribution. Environment. The climate in the 64.03 Hawkdun and 64.04 Saint Bathans Ecological Districts is semi-arid with warm, dry summers and cold winters (McEwen 1987). LENZ Level IV data indicate the land environment at the type locality is particularly harsh, with an annual average temperature of 4.2 °C, a mean minimum temperature of -4.1 °C during the coldest month of the austral winter (July), average annual solar radiation of 13.2 MJ/m2/day and winter solar radiation of 3.8 MJ/m2 /day (Leathwick et al. 2002). Habitat. Fractured and deeply creviced greywacke outcrops and deep boulderfields and talus slopes. Snow tussock (Chionochloa rigida) grassland and Dracophyllum rosmarinifolium shrubland are the predominant vegetation types in the areas where the species has been found, interrupted by greywacke rock outcrops or boulderfields and talus. At the type locality, M. galaxias have been found on six discrete, but proximate rock outcrops and bluffs comprised of a protruding but deeply fissured and fractured central tor several metres tall, surrounded by varying amounts of rock talus, fellfield and/or scree, along a ~270-m section of a ridgeline. The first gecko discovered was located within a crevice inside a c. 50 cm fractured rock, in an expanse of stony ground c. 10 m from a major outcrop system. The other eighteen geckos encountered at the type locality have all been inside vertical rock fissures, or out on rock ledges. At the other three sites (across two mountain ranges) the five geckos observed were emergent in deep boulderfield or talus, but three of these were well concealed by surrounding rocks and were in positions close to retreats or escape routes. Some of the rocks at all known sites, particularly on tors, are heavily encrusted with colourful lichens of various species, including large amounts of the tall, bristly Neuropogon spp., the latter which may limit the extent of usable rock surface for the geckos (Fig. 6). Natural history. Mokopirirakau galaxias is apparently an alpine specialist gecko, saxicolous in greywacke rock outcrops, boulderfields, and talus. The geckos are strongly nocturnal and thigmothermic. They have been detected on exposed rock surfaces during surveys at night in air temperatures of 9-16 °C. Nothing is known of the species’ life history, diet or reproduction. The sex ratio appears to be close to 1:1 based on the limited survey results to date (C. Knox, unpub. data). A small number of juvenile and subadult geckos have been located, but not neonates. It is likely to be mostly insectivorous, but may consume berries of alpine shrubs where available. It is likely to be viviparous, as all New Zealand geckos are. However, due to the species’ alpine niche, it is most likely slow-growing, late to mature (at least 5 years) and long lived (to decadal timeframes), with a very low reproductive output (Cree 1994, Cree & Hare 2016; Knox et al. 2019). One of two adult females captured in January 2018 and a female captured in November 2020 were pregnant; however, no inference can be made on the annual reproductive output per female based on this very small sample size. At all known sites M. galaxias is sympatric with the Southern Alps gecko (Woodworthia “Southern Alps”), which is sometimes numerous and occupies all available rocky habitat from outcrops to talus and scree. The two gecko species can be syntopic, sharing the same rock crevice or foraging on the same rock ledge. Size differences between M. galaxias (to 84 mm SVL) and W. “Southern Alps” (c. 55-65 mm SVL) may allow resource partitioning in retreat site and prey selection between the two species. In contrast, Whitaker et al. (1999) noted that in Nelson and Marlborough, M. kahutarae appear to exclude specimens of the sympatric Woodworthia geckos from their immediate territory. Several species of Oligosoma skinks are also present, particularly O. maccanni, which are found throughout the same rocky habitat and in the surrounding alpine tussockland. Phylogenetic affinities. Genetic analyses using sequences from the mitochondrial ND2 gene indicate that M. galaxias is sister to a clade comprising the sister species M. granulatus and M. kahutarae. Mokopirirakau granulatus occurs in the upper half of the North Island, and in Marlborough, Tasman, and Westland in the South Island. The closest M. kahutarae populations to M. galaxias are approximately 400 km to the north-east; whereas, the closest M. granulatus populations to M. galaxias are approximately 180 km away, but on the other side of the Southern Alps (Fig. 7). Thus, M. galaxias is completely isolated from its nearest relatives geographically, with multiple mountain ranges between their known ranges. The geographically closest Mokopirirakau populations are of the undescribed taxon M. “Roy’s Peak”, which occurs to the west and south-west (van Winkel et al. 2018; Knox et al. 2019). Conservation. At the type locality this species is currently known only from eight rock tors and surrounding talus within a 1.4-ha area, and 10 °C, dry and calm nights), but did not locate new populations. However, in November and December 2020, three new populations were found, including two sites in another mountain range within Oteake Conservation Park. Given the November and December 2020 finds, it is deemed likely that other populations of this species will be discovered, as further survey work is undertaken in North Otago and South Canterbury. Threats to M. galaxias include predation from introduced mammals, illegal collection, and climate change. In addition, fires and grazing by introduced herbivores may be a threat by reducing habitat quality and food availability. Introduced mammalian carnivores and rodents that may encroach into their high altitude, exposed habitat pose a threat, especially predators that hunt at night when the geckos may be emergent on rock surfaces. These are likely to include cats (Felix catus), ferrets (Mustela furo), stoats (Mustela erminea), weasels (Mustela nivalis), hedgehogs (Erinaceus europaeus), possums (Trichosurus vulpecula), and mice (Mus musculus) (O’Donnell et al. 2017; Josh Kemp pers. comm.). Mice are likely to have population irruptions following tussock mast seeding, which will then drive secondary carnivore irruptions. Stoats are known to prey on alpine lizards (O’Donnell et al. 2017), and mice and weasels can squeeze into tight crevices to access the geckos inside retreats. Illegal collection has emerged as a threat to several species of New Zealand lizards, particularly geckos (Knox et al. 2013; Marshall et al. 2020) and if precise site information becomes publicly available then determined smugglers could impact the abundance of geckos at those sites. Climate change may also render any remaining habitat increasingly vulnerable to introduced mammals, as it is expected that more predators will infiltrate alpine areas as the climate warms (O’Donnell et al. 2017). Due to the species being discovered very recently, it is not yet listed in the most recent iteration of the Department of Conservation’s New Zealand Threat Classification System lists (Hitchmough et al. 2016). Typically, newly discovered species are placed in the “Data Deficient” category. However, the system requires a precautionary ap- proach under which, provided some information exists, the species is placed into the most threatened category consistent with what is known, with uncertainty being indicated by a “Data Poor” qualifier. We recommend the species be assigned the conservation ranking of “Threatened—Nationally Endangered” with the qualifier “Data Poor” (Townsend et al. 2008). This is because of its low abundance, likely decline resulting from impacts of predatory mammals, threats from climate change, and highly restricted range (four small locations). This species should be a high priority for the Department of Conservation for surveys to improve understanding of its distribution, conservation status and conservation management requirements. In the IUCN Red List system, in the absence of clear information on population trend, Data Deficient seems the best recommendation for listing.

Naultinus flavirictus sp. nov. (Fig. 1) Naultinus grayi: (part) Robb & Hitchmough (1980). Naultinus “North Cape”: Hitchmough et al. (2007). Holotype. Adult female holotype MONZ RE003322, original field number G.789 (Fig. 1), collected from Te Paki, Northland, New Zealand, collected by A. H. Whitaker in May 1972. Paratypes. Two adults, one male, MONZ RE.005611 (original field number RAH195), collected from the Te Kao District, Northland, New Zealand, by Conrad Hepers in August 1986, and one female, MONZ RE.005609 (original field number RAH254; captive offspring of a female collected from Te Kao, Northland, New Zealand, by Conrad Hepers in August 1986). Etymology. Naultinus flavirictus sp. nov. (formed from the Lain flavus = yellow + rictus = gaping of the mouth) is named for the continuation of the yellow-orange colouring of the jaws and tongue onto the external corners of the mouth, which is diagnostic for the species. The epithet is formed as a noun in apposition. Diagnosis. Naultinus flavirictus sp. nov. is a small Naultinus (maximum SVL ~ 70 mm), with a short, deep snout, fine granular body scales, flattened scales on the dorsum of the snout, rostral scale divided by a median crease, pale lavender mouth lining, and yellow-orange jaws, tongue and external corners of the mouth. It may be distinguished from all other North Island Naultinus species by the pale lavender mouth lining, with yellow-orange jaws and tongue, and by the colour of the jaws extending to the external corners of the mouth, with similar pigment present around the nostrils. A central crease dividing rostral in two is typical in this species, but rare in N. grayii and N. elegans, and partial or absent in other Naultinus. It is distinguished from N. grayii and N. punctatus by much smaller adult size (to ~ 70 mm vs ~ 95 mm SVL), and by secondary bluish colouration of adult males including the entire ventral surface rather than being confined to a lower lateral band. It is further distinguished from N. grayii by its proportionally shorter, deeper snout (Fig. 2). Like N. grayii, but unlike all other Naultinus, the scales on the dorsal surface of the snout are flattened (Fig. 2). The new species may be distinguished from most South Island Naultinus by its fine, granular dorsal body scales (coarser and more conical in South Island species). Among its South Island congeners it is readily distinguished from N. rudis (Fischer), N. tuberculatus and Otago / Southland populations of N. gemmeus by the pale lavender and orange mouth lining. It differs from N. manukanus and N. rudis in the absence of any greatly enlarged dorsal body and head scales (although there is a north–south cline in the number and prominence of these enlarged scales within N. manukanus, and they may be few in number and easily missed in the northernmost population on Stephens Island; this population is morphologically similar to north-western-most populations of N. stellatus) (Bull & Whitaker 1975). It may be distinguished from Canterbury populations of N. gemmeus by its smaller adult size, and by males having bluish belly colouration rather than having the green colouration progressively replaced by brown or grey during growth to maturity. It is most similar morphologically to north-western-most populations of N. stellatus, but distinguished by more subimbricate, slightly flattened granular (vs strongly domed, bead-like, tightly juxtaposed) dorsal body scales, less domed canthal scales (Fig. 2), presence of an enlarged patch of supratemporal scales, and rows of enlarged, equal-sized scales each side of the canthal row in N. flavirictus sp. nov. (vs irregular-sized scales bordering enlarged canthal row in N. stellatus). Naultinus flavirictus sp. nov. is distinguished from other populations of N. stellatus by its uniform green background colour, with or without simple, discrete, irregular-shaped spotted or striped markings rather than large, circular blotches with concentric rings of colour. Description of the holotype. Body moderately slender, not depressed; 67.1 mm SVL. HeadL 16.6 mm (25% SVL), HeadW 12.1 mm (73% HeadL) and HeadH 8.8 mm (53% HL). Head subtriangular with a blunt, rounded snout, clearly demarcated from neck; interorbital/frontal region neither concave nor convex, nasal region without indentation; canthus not inflated. SnEye 7.5 mm (45% HeadL), OrbD 3.4 mm (45% SnEye). Scales on dorsum of anterior snout approximately 3-4 times the diameter of those on occipital region, loreal scales similar, abruptly decreasing in size at level of anterior edge of orbit; patches of enlarged supratemporal scales (smaller than dorsal snout and loreal scales but distinctly larger than occipital granules). OrbD 20% HeadL; pupil oval, margins smooth. Ear opening very small (1.1 mm), oval, obliquely oriented; eye to ear distance greater than diameter of eyes (EyeEar 5.6 mm (165% OrbD). Rostral almost rectangular, much broader (3.0 mm) than high (1.3 mm) completely divided by a strong rostral crease, contacted dorsally by one large round internasal and two large supranasals, each similar in size to the internasal, contacted laterally by first supralabial, excluded from nostril. Nostrils rounded, laterally oriented, and surrounded by two postnasals on right hand side, three on left, first supralabial and first supranasal. Mental subtriangular, broader (2.8 mm) than tall (1.6 mm). First infralabials separated by three postmental, the central one largest and oval, longer than wide, the others wider than long. Anterior infralabials and postmental bordered posteriorly by 3–4 series of enlarged hexagonal chin shields, ¼ to ½ the size of central postmental chin shield, changing abruptly to small, rounded scales beneath throat. Scale rows ventral to posterior infralabials on lateral surface of jaw slightly enlarged and elongated relative to ventral throat scales. Eleven (right) and 10 (left) enlarged infralabial scales, posterior most about 4 times size of rictal scales, 10 (left)–12 (right) enlarged supralabial scales, 7 (left) or 10 (right) supralabials to midpoint of orbit; 36 scale rows between supraciliaries (24 between supraocular skin folds), 11 scale rows across frontal bones at midpoint of orbit. Supraciliaries rounded to somewhat conical posteriorly. Dorsal scales small, mostly uniform in size with scattered much smaller scales interspersed, homogenous in shape, bead-like, somewhat flattened, granular. Ventral scales roughly twice diameter of dorsals, smooth, flattened, subimbricate, slightly enlarged in precloacal region. Approximately 99 scale rows around mid-body. No ventrolateral skin folds. Scales on dorsal surface of upper forelimb roughly equal in size to dorsum; on forearm slightly smaller. Scales on preaxial surface of thigh similar to those on dorsum, gradually decreasing in size posteriorly to smaller granules on post-axial surface. Scales on palms and soles smooth, flattened and subimbricate. Fore- and hindlimbs short and moderately robust (Forearm 8.3 mm; ForeaL/SVL 0.12; Crus 11.1 mm; CrusL/SVL 0.17). Digits relatively elongate, all bearing claws. Claws short and moderately robust basally, recurved to a fine point. Basal portion of digits relatively narrow, approximately twice width of distal phalanges. Distal portion of toe strongly compressed, lamellae continuing in uninterrupted series to claw. Apical pads absent. Relative length of digits of manus: I twice size of scales at base of digits) from right (and left) sides 7-10-13-14-11 (6-9-14-14-11) manus and 7-13-14-16-15 (7-13-15-17-15) pes. Only scattered pitted scales present in precloacal area (c. 5 rows). Small, weakly developed cloacal spurs with 3 enlarged scales on each side just posterior to vent. Tail original but broken (9.5 +79.7 = 89.2 mm); 133% snoutvent length, muscular, tapering, round in cross-section. Dorsal caudal scales medium, flat, juxtaposed to weakly subimbricate, roughly circular, arranged irregularly. Surface of tail not segmented. Ventral scales slightly larger than dorsal, smooth, somewhat irregular in shape, imbricate. Colour in preservative. Dorsum uniform dull dark green with dorso-lateral rows of irregularly shaped pale cream spots, comprising paired longitudinal streaks behind the eyes, seven spots on each side between the neck and the pelvic region; spots continue along the tail where they become elongated into streaks. Uniform whitish ventro-lateral stripes between insertions of fore and hind limbs separating dark green lateral from pale green ventral surface. Small white blotch of c. 7 scales on dorsum of left thigh. Lower lip with strong white marking extending almost to earhole (Figs. 1, 2). Faint yellow colouration around nostrils. Undersides of feet and toes similar in colouring to venter. Colour in life. (from paratypes, supplemented by non-vouchered photographs of other wild individuals – Fig. 3). As in preservative, except that the basic colour is very bright green rather than dull dark green. Pattern of spots or stripes is variable among individuals, or can be absent. There is sexual dichromatism, with adult and subadult males having the ventral surface and lower lateral surface very pale blue; these areas are light green in females. In a few individuals the background bright green colouration is replaced by bright yellow (P. Miller pers. comm.; van Winkel et al. 2018). Pale lavender mouth lining, with yellow-orange jaws and tongue, with the colour of the jaws extending to the external corners of the mouth, and with similar pigment present around the nostrils. Paratype variation. Both MONZ RE.005611 and MONZ RE.005609 are very similar in appearance to the holotype. They differ in the extent of pale dorsal markings, and presence of mid-lateral markings in MONZ RE.005611), number and arrangement of internasals and postmentals, and have minor differences in numbers of infralabials and supralabials, scale rows between supraciliaries, scale rows across frontal bones at midpoint of orbit, lamellae counts and numbers of precloacal pores (which are more strongly developed in the male paratype). MONZ RE.005611 has a small, diamond-shaped internasal, posterior to which are a pair of considerably larger, hexagonal internasals, which exclude the small central one from contact with the supranasals. Four postnasals on left, three on right. Two hexagonal postmentals. Chin shields ¼ to same size as postmentals. 11 (left), 10 (right) infralabials. 13 (left), 11 (right) supralabials. 9th (left), 8th (right) in mid-orbital position. 39 scale rows between supraciliaries (29 between supraocular skin folds), 10 scale rows across frontal bones at midpoint of orbit. Dorsal and particularly lateral scales weakly subimbricate. Lamellar counts (including all scales> twice size of scales at base of digits) from right (and left) sides 5-12-13-14-10 (6-11-13-14-12) manus and 7-13-15-14-15 (7-13-14-14-13) pes. Precloacal pores 112 in total, in 5 rows, 3 extending onto thigh, 2 reaching mid-point of thigh. Scale row immediately posterior to precloacal spores includes 5 pitted scales – not included in count. Four left and three right, enlarged, raised and sharply pointed cloacal spurs. Colour in preservative deep lavender dorsally, becoming paler on flanks and limbs, body with paired, mostly symmetrical series of cream-coloured markings, fragmented into several elongate lines on left side, series of more variable lines and blotches on right side, extending onto tail as two broken lines; head with a pair of crescentic supratemporal markings, a large marking extending between left and right antorbital positions, narrowing in the interorbital region and expanding again into a blunted arrowhead shape on the parietals. All except last marking outlined by a thin, more or less continuous black line. Narrow cream line extending from infralabials almost to ear opening. Ventrolateral marking as in holotype. Small scattered white patches on posterior flanks (one on left, two on right). Limbs without markings. Venter immaculate. Dorsum and venter of digits pale straw colour. MONZ RE.005609 has a single hexagonal internasal and three postnasals on both left and right sides. Two irregular postmentals, right three times size of left. Chin shields 1/3 to same size as postmentals. 11 (left), 12 (right) infralabials. 13 (left), 14 (right) supralabials. 10th (left), 11th (right) in mid-orbital position. 37 scale rows between supraciliaries (27 between supraocular skin folds), 11 scale rows across frontal bones at midpoint of orbit. Dorsal and particularly lateral scales weakly subimbricate. Lamellar counts (including all scales> twice size of scales at base of digits) from right (and left) sides 5 (other digits damaged) (5-9-10-11-10) manus and 5-12- damaged-damaged-14 (6-damaged-damaged-14-damaged) pes. Six rows of pitted scales (c. 75 total) only one extending onto base of thigh. Four left and three right, enlarged, raised and sharply pointed cloacal spurs. Colour in preservative pale lavender, only visible dorsal markings are faded, slightly bowed, whitish supratemporal markings. Geographic distribution. The northern half of the Aupōuri Peninsula, the peninsula at the northern extremity of the North Island of New Zealand. Most records are from the Te Paki area in the far north (a general distribution map is provided by Nielsen et al. 2011). The gecko is found on the northern half of the peninsula, from about Houhora northwards. Exact locality details are not provided to discourage illegal collection, which is a well-documented threat to New Zealand’s native herpetofauna (Nelson et al. 2014; Hitchmough et al. 2016) and globally (Bryan et al. 2006). Biogeography. The paleo-islands at the northern end of the peninsula are formed of Cretaceous ophiolitic (basalt, gabbro and serpentinite) and Lower Miocene marine sedimentary rocks (Brook & Thrasher 1991). These paleo-islands were probably emergent above sea-level from Late Miocene or Early Pliocene time. During Early to Mid Pleistocene time extensive deposits of marine and aeolian sand accumulated north of Kaitaia; paleo-islands probably existed in the Reinga-North Cape and Houhora areas during interglacial periods (i.e., high sea levels), but a proto-Aupouri Peninsula would have existed during glacial low stands. Dune buildup during Late Pleistocene time formed the present land connection along the Peninsula. Thus, the far northern islands were probably isolated throughout the Pliocene, and were intermittently connected with the rest of Northland during the Pleistocene, with the connection becoming permanent in the Late Pleistocene (Brook & Thrasher 1991). Although we did not perform specific dating analyses per se, if we assume a mutation rate for ND2 of 0.65% per million years (as proposed by Macey et al. 1998), the ~4% mitochondrial distance separating N. flavirictus sp. nov. from its closest kin (Table S01) correlates to ~2.5 million years. It is therefore reasonable to infer that the ancestors of Naultinus flavirictus sp. nov. reached the Te Paki area while it was an island or series of islands and evolved there in isolation until the formation of the Aupōuri tombolo enabled it to come back into contact with N. grayii during the Pleistocene. Contact zones with N. grayii may have repeatedly formed and then been removed by inter-glacial sea-level rise, and the current contact is likely to have formed in the Late Pleistocene. Reproduction. Animals collected from Te Kao in August 1986 then held in captivity in Auckland gave birth to twins in late January 1987 (C. Hepers, pers. comm.). This is almost two months earlier than births of N. grayii, both in the field and in captivity in Auckland (Robb & Hitchmough 1980). Conservation status. This species is listed by Hitchmough et al. (2016) under the New Zealand Threat Classification System as “At Risk – Declining” on the basis of criterion C (2/1) (total area of occupancy> 100 km 2, predicted decline 10–70%), and with the Qualifiers Data Poor and One Location. The One Location qualifier means that the species occupies a single area of continuous habitat, so a new threat could very quickly spread to affect the entire population. Under the IUCN Redlist system, we suggest the species is Endangered under the criteria B1a,b(v), B2a,b(v): Geographic range in the form of (B1) extent of occurrence estimated to be (1) less than 5,000 km 2, and (a) estimates indicating severely fragmented or known to exist at no more than five locations and (b) continuing decline, observed, inferred or projected, in (v) number of mature individuals. Geographic range in the form of (B2) area of occupancy estimated to be (1) less than 500 km 2, and estimates indicating (a) severely fragmented or known to exist at no more than five locations and (b) continuing decline, observed, inferred or projected, in (v) number of mature individuals. The presumed original habitat of N. flavirictus sp. nov., tall subtropical rainforest, has been almost entirely removed since human arrival in the 13 th Century AD, and now only exists in a handful of tiny remnant patches (3.2% of the Ecological District; Lux et al. 2009). Fires set by Māori removed most of the original forest on the Aupōuri Peninsula soon after their arrival in New Zealand in about the 12 th Century CE; initially it was replaced by extensive areas of bracken fern (Pteridium esculentum (G. Forster)) (Elliot et al. 1995). The Te Paki area is now largely covered in fire-maintained secondary shrublands dominated by Kunzea Reichenbach and Leptospermum J.R. Forster & G. Forster. However, this shrubland is readily occupied by Naultinus, so the total area of habitat has probably changed relatively little. Only parts of the available habitat are known to be occupied, probably as a result of fire history. With fire now better controlled, threats are almost entirely from invasive species. Almost the full suite of small predatory introduced mammals which have decimated native lizard populations across mainland New Zealand is present and common: cats (Felis catus Linnaeus), hedgehogs (Erinaceus europaeus Linnaeus), brush-tailed possums (Trichosurus vulpecula (Kerr)), stoats (Mustela erminea Linnaeus), weasels (Mustela nivalis Linnaeus), mice (Mus musculus Linnaeus), and rats (both Rattus rattus (Linnaeus) and R. norvegicus (Berkenhout)). Feral pigs (Sus scrofa Linnaeus), which have also recently been demonstrated to eat large numbers of native lizards (J. Reardon pers. comm.), are widespread and abundant. Te Paki was one of the last areas of the country reached by possums, which have become abundant only in the last 20 years. In 1985 they were still recorded as absent but were spreading rapidly northward up the Aupōuri Peninsula, where they had reached about 10 km south of Waitiki Landing (Mitchell 1985). The relatively abundant (c

Naultinus flavirictus sp. nov. (Fig. 1) Naultinus grayi: (part) Robb & Hitchmough (1980). Naultinus ���North Cape���: Hitchmough et al. (2007). Holotype. Adult female holotype MONZ RE003322, original field number G.789 (Fig. 1), collected from Te Paki, Northland, New Zealand, collected by A. H. Whitaker in May 1972. Paratypes. Two adults, one male, MONZ RE.005611 (original field number RAH195), collected from the Te Kao District, Northland, New Zealand, by Conrad Hepers in August 1986, and one female, MONZ RE.005609 (original field number RAH254; captive offspring of a female collected from Te Kao, Northland, New Zealand, by Conrad Hepers in August 1986). Etymology. Naultinus flavirictus sp. nov. (formed from the Lain flavus = yellow + rictus = gaping of the mouth) is named for the continuation of the yellow-orange colouring of the jaws and tongue onto the external corners of the mouth, which is diagnostic for the species. The epithet is formed as a noun in apposition. Diagnosis. Naultinus flavirictus sp. nov. is a small Naultinus (maximum SVL ~ 70 mm), with a short, deep snout, fine granular body scales, flattened scales on the dorsum of the snout, rostral scale divided by a median crease, pale lavender mouth lining, and yellow-orange jaws, tongue and external corners of the mouth. It may be distinguished from all other North Island Naultinus species by the pale lavender mouth lining, with yellow-orange jaws and tongue, and by the colour of the jaws extending to the external corners of the mouth, with similar pigment present around the nostrils. A central crease dividing rostral in two is typical in this species, but rare in N. grayii and N. elegans, and partial or absent in other Naultinus. It is distinguished from N. grayii and N. punctatus by much smaller adult size (to ~ 70 mm vs ~ 95 mm SVL), and by secondary bluish colouration of adult males including the entire ventral surface rather than being confined to a lower lateral band. It is further distinguished from N. grayii by its proportionally shorter, deeper snout (Fig. 2). Like N. grayii, but unlike all other Naultinus, the scales on the dorsal surface of the snout are flattened (Fig. 2). The new species may be distinguished from most South Island Naultinus by its fine, granular dorsal body scales (coarser and more conical in South Island species). Among its South Island congeners it is readily distinguished from N. rudis (Fischer), N. tuberculatus and Otago / Southland populations of N. gemmeus by the pale lavender and orange mouth lining. It differs from N. manukanus and N. rudis in the absence of any greatly enlarged dorsal body and head scales (although there is a north���south cline in the number and prominence of these enlarged scales within N. manukanus, and they may be few in number and easily missed in the northernmost population on Stephens Island; this population is morphologically similar to north-western-most populations of N. stellatus) (Bull & Whitaker 1975). It may be distinguished from Canterbury populations of N. gemmeus by its smaller adult size, and by males having bluish belly colouration rather than having the green colouration progressively replaced by brown or grey during growth to maturity. It is most similar morphologically to north-western-most populations of N. stellatus, but distinguished by more subimbricate, slightly flattened granular (vs strongly domed, bead-like, tightly juxtaposed) dorsal body scales, less domed canthal scales (Fig. 2), presence of an enlarged patch of supratemporal scales, and rows of enlarged, equal-sized scales each side of the canthal row in N. flavirictus sp. nov. (vs irregular-sized scales bordering enlarged canthal row in N. stellatus). Naultinus flavirictus sp. nov. is distinguished from other populations of N. stellatus by its uniform green background colour, with or without simple, discrete, irregular-shaped spotted or striped markings rather than large, circular blotches with concentric rings of colour. Description of the holotype. Body moderately slender, not depressed; 67.1 mm SVL. HeadL 16.6 mm (25% SVL), HeadW 12.1 mm (73% HeadL) and HeadH 8.8 mm (53% HL). Head subtriangular with a blunt, rounded snout, clearly demarcated from neck; interorbital/frontal region neither concave nor convex, nasal region without indentation; canthus not inflated. SnEye 7.5 mm (45% HeadL), OrbD 3.4 mm (45% SnEye). Scales on dorsum of anterior snout approximately 3-4 times the diameter of those on occipital region, loreal scales similar, abruptly decreasing in size at level of anterior edge of orbit; patches of enlarged supratemporal scales (smaller than dorsal snout and loreal scales but distinctly larger than occipital granules). OrbD 20% HeadL; pupil oval, margins smooth. Ear opening very small (1.1 mm), oval, obliquely oriented; eye to ear distance greater than diameter of eyes (EyeEar 5.6 mm (165% OrbD). Rostral almost rectangular, much broader (3.0 mm) than high (1.3 mm) completely divided by a strong rostral crease, contacted dorsally by one large round internasal and two large supranasals, each similar in size to the internasal, contacted laterally by first supralabial, excluded from nostril. Nostrils rounded, laterally oriented, and surrounded by two postnasals on right hand side, three on left, first supralabial and first supranasal. Mental subtriangular, broader (2.8 mm) than tall (1.6 mm). First infralabials separated by three postmental, the central one largest and oval, longer than wide, the others wider than long. Anterior infralabials and postmental bordered posteriorly by 3���4 series of enlarged hexagonal chin shields, �� to �� the size of central postmental chin shield, changing abruptly to small, rounded scales beneath throat. Scale rows ventral to posterior infralabials on lateral surface of jaw slightly enlarged and elongated relative to ventral throat scales. Eleven (right) and 10 (left) enlarged infralabial scales, posterior most about 4 times size of rictal scales, 10 (left)���12 (right) enlarged supralabial scales, 7 (left) or 10 (right) supralabials to midpoint of orbit; 36 scale rows between supraciliaries (24 between supraocular skin folds), 11 scale rows across frontal bones at midpoint of orbit. Supraciliaries rounded to somewhat conical posteriorly. Dorsal scales small, mostly uniform in size with scattered much smaller scales interspersed, homogenous in shape, bead-like, somewhat flattened, granular. Ventral scales roughly twice diameter of dorsals, smooth, flattened, subimbricate, slightly enlarged in precloacal region. Approximately 99 scale rows around mid-body. No ventrolateral skin folds. Scales on dorsal surface of upper forelimb roughly equal in size to dorsum; on forearm slightly smaller. Scales on preaxial surface of thigh similar to those on dorsum, gradually decreasing in size posteriorly to smaller granules on post-axial surface. Scales on palms and soles smooth, flattened and subimbricate. Fore- and hindlimbs short and moderately robust (Forearm 8.3 mm; ForeaL/SVL 0.12; Crus 11.1 mm; CrusL/SVL 0.17). Digits relatively elongate, all bearing claws. Claws short and moderately robust basally, recurved to a fine point. Basal portion of digits relatively narrow, approximately twice width of distal phalanges. Distal portion of toe strongly compressed, lamellae continuing in uninterrupted series to claw. Apical pads absent. Relative length of digits of manus: I twice size of scales at base of digits) from right (and left) sides 7-10-13-14-11 (6-9-14-14-11) manus and 7-13-14-16-15 (7-13-15-17-15) pes. Only scattered pitted scales present in precloacal area (c. 5 rows). Small, weakly developed cloacal spurs with 3 enlarged scales on each side just posterior to vent. Tail original but broken (9.5 +79.7 = 89.2 mm); 133% snoutvent length, muscular, tapering, round in cross-section. Dorsal caudal scales medium, flat, juxtaposed to weakly subimbricate, roughly circular, arranged irregularly. Surface of tail not segmented. Ventral scales slightly larger than dorsal, smooth, somewhat irregular in shape, imbricate. Colour in preservative. Dorsum uniform dull dark green with dorso-lateral rows of irregularly shaped pale cream spots, comprising paired longitudinal streaks behind the eyes, seven spots on each side between the neck and the pelvic region; spots continue along the tail where they become elongated into streaks. Uniform whitish ventro-lateral stripes between insertions of fore and hind limbs separating dark green lateral from pale green ventral surface. Small white blotch of c. 7 scales on dorsum of left thigh. Lower lip with strong white marking extending almost to earhole (Figs. 1, 2). Faint yellow colouration around nostrils. Undersides of feet and toes similar in colouring to venter. Colour in life. (from paratypes, supplemented by non-vouchered photographs of other wild individuals ��� Fig. 3). As in preservative, except that the basic colour is very bright green rather than dull dark green. Pattern of spots or stripes is variable among individuals, or can be absent. There is sexual dichromatism, with adult and subadult males having the ventral surface and lower lateral surface very pale blue; these areas are light green in females. In a few individuals the background bright green colouration is replaced by bright yellow (P. Miller pers. comm.; van Winkel et al. 2018). Pale lavender mouth lining, with yellow-orange jaws and tongue, with the colour of the jaws extending to the external corners of the mouth, and with similar pigment present around the nostrils. Paratype variation. Both MONZ RE.005611 and MONZ RE.005609 are very similar in appearance to the holotype. They differ in the extent of pale dorsal markings, and presence of mid-lateral markings in MONZ RE.005611), number and arrangement of internasals and postmentals, and have minor differences in numbers of infralabials and supralabials, scale rows between supraciliaries, scale rows across frontal bones at midpoint of orbit, lamellae counts and numbers of precloacal pores (which are more strongly developed in the male paratype). MONZ RE.005611 has a small, diamond-shaped internasal, posterior to which are a pair of considerably larger, hexagonal internasals, which exclude the small central one from contact with the supranasals. Four postnasals on left, three on right. Two hexagonal postmentals. Chin shields �� to same size as postmentals. 11 (left), 10 (right) infralabials. 13 (left), 11 (right) supralabials. 9th (left), 8th (right) in mid-orbital position. 39 scale rows between supraciliaries (29 between supraocular skin folds), 10 scale rows across frontal bones at midpoint of orbit. Dorsal and particularly lateral scales weakly subimbricate. Lamellar counts (including all scales> twice size of scales at base of digits) from right (and left) sides 5-12-13-14-10 (6-11-13-14-12) manus and 7-13-15-14-15 (7-13-14-14-13) pes. Precloacal pores 112 in total, in 5 rows, 3 extending onto thigh, 2 reaching mid-point of thigh. Scale row immediately posterior to precloacal spores includes 5 pitted scales ��� not included in count. Four left and three right, enlarged, raised and sharply pointed cloacal spurs. Colour in preservative deep lavender dorsally, becoming paler on flanks and limbs, body with paired, mostly symmetrical series of cream-coloured markings, fragmented into several elongate lines on left side, series of more variable lines and blotches on right side, extending onto tail as two broken lines; head with a pair of crescentic supratemporal markings, a large marking extending between left and right antorbital positions, narrowing in the interorbital region and expanding again into a blunted arrowhead shape on the parietals. All except last marking outlined by a thin, more or less continuous black line. Narrow cream line extending from infralabials almost to ear opening. Ventrolateral marking as in holotype. Small scattered white patches on posterior flanks (one on left, two on right). Limbs without markings. Venter immaculate. Dorsum and venter of digits pale straw colour. MONZ RE.005609 has a single hexagonal internasal and three postnasals on both left and right sides. Two irregular postmentals, right three times size of left. Chin shields 1/3 to same size as postmentals. 11 (left), 12 (right) infralabials. 13 (left), 14 (right) supralabials. 10th (left), 11th (right) in mid-orbital position. 37 scale rows between supraciliaries (27 between supraocular skin folds), 11 scale rows across frontal bones at midpoint of orbit. Dorsal and particularly lateral scales weakly subimbricate. Lamellar counts (including all scales> twice size of scales at base of digits) from right (and left) sides 5 (other digits damaged) (5-9-10-11-10) manus and 5-12- damaged-damaged-14 (6-damaged-damaged-14-damaged) pes. Six rows of pitted scales (c. 75 total) only one extending onto base of thigh. Four left and three right, enlarged, raised and sharply pointed cloacal spurs. Colour in preservative pale lavender, only visible dorsal markings are faded, slightly bowed, whitish supratemporal markings. Geographic distribution. The northern half of the Aupōuri Peninsula, the peninsula at the northern extremity of the North Island of New Zealand. Most records are from the Te Paki area in the far north (a general distribution map is provided by Nielsen et al. 2011). The gecko is found on the northern half of the peninsula, from about Houhora northwards. Exact locality details are not provided to discourage illegal collection, which is a well-documented threat to New Zealand���s native herpetofauna (Nelson et al. 2014; Hitchmough et al. 2016) and globally (Bryan et al. 2006). Biogeography. The paleo-islands at the northern end of the peninsula are formed of Cretaceous ophiolitic (basalt, gabbro and serpentinite) and Lower Miocene marine sedimentary rocks (Brook & Thrasher 1991). These paleo-islands were probably emergent above sea-level from Late Miocene or Early Pliocene time. During Early to Mid Pleistocene time extensive deposits of marine and aeolian sand accumulated north of Kaitaia; paleo-islands probably existed in the Reinga-North Cape and Houhora areas during interglacial periods (i.e., high sea levels), but a proto-Aupouri Peninsula would have existed during glacial low stands. Dune buildup during Late Pleistocene time formed the present land connection along the Peninsula. Thus, the far northern islands were probably isolated throughout the Pliocene, and were intermittently connected with the rest of Northland during the Pleistocene, with the connection becoming permanent in the Late Pleistocene (Brook & Thrasher 1991). Although we did not perform specific dating analyses per se, if we assume a mutation rate for ND2 of 0.65% per million years (as proposed by Macey et al. 1998), the ~4% mitochondrial distance separating N. flavirictus sp. nov. from its closest kin (Table S01) correlates to ~2.5 million years. It is therefore reasonable to infer that the ancestors of Naultinus flavirictus sp. nov. reached the Te Paki area while it was an island or series of islands and evolved there in isolation until the formation of the Aupōuri tombolo enabled it to come back into contact with N. grayii during the Pleistocene. Contact zones with N. grayii may have repeatedly formed and then been removed by inter-glacial sea-level rise, and the current contact is likely to have formed in the Late Pleistocene. Reproduction. Animals collected from Te Kao in August 1986 then held in captivity in Auckland gave birth to twins in late January 1987 (C. Hepers, pers. comm.). This is almost two months earlier than births of N. grayii, both in the field and in captivity in Auckland (Robb & Hitchmough 1980). Conservation status. This species is listed by Hitchmough et al. (2016) under the New Zealand Threat Classification System as ���At Risk ��� Declining��� on the basis of criterion C (2/1) (total area of occupancy> 100 km 2, predicted decline 10���70%), and with the Qualifiers Data Poor and One Location. The One Location qualifier means that the species occupies a single area of continuous habitat, so a new threat could very quickly spread to affect the entire population. Under the IUCN Redlist system, we suggest the species is Endangered under the criteria B1a,b(v), B2a,b(v): Geographic range in the form of (B1) extent of occurrence estimated to be (1) less than 5,000 km 2, and (a) estimates indicating severely fragmented or known to exist at no more than five locations and (b) continuing decline, observed, inferred or projected, in (v) number of mature individuals. Geographic range in the form of (B2) area of occupancy estimated to be (1) less than 500 km 2, and estimates indicating (a) severely fragmented or known to exist at no more than five locations and (b) continuing decline, observed, inferred or projected, in (v) number of mature individuals. The presumed original habitat of N. flavirictus sp. nov., tall subtropical rainforest, has been almost entirely removed since human arrival in the 13 th Century AD, and now only exists in a handful of tiny remnant patches (3.2% of the Ecological District; Lux et al. 2009). Fires set by Māori removed most of the original forest on the Aupōuri Peninsula soon after their arrival in New Zealand in about the 12 th Century CE; initially it was replaced by extensive areas of bracken fern (Pteridium esculentum (G. Forster)) (Elliot et al. 1995). The Te Paki area is now largely covered in fire-maintained secondary shrublands dominated by Kunzea Reichenbach and Leptospermum J.R. Forster & G. Forster. However, this shrubland is readily occupied by Naultinus, so the total area of habitat has probably changed relatively little. Only parts of the available habitat are known to be occupied, probably as a result of fire history. With fire now better controlled, threats are almost entirely from invasive species. Almost the full suite of small predatory introduced mammals which have decimated native lizard populations across mainland New Zealand is present and common: cats (Felis catus Linnaeus), hedgehogs (Erinaceus europaeus Linnaeus), brush-tailed possums (Trichosurus vulpecula (Kerr)), stoats (Mustela erminea Linnaeus), weasels (Mustela nivalis Linnaeus), mice (Mus musculus Linnaeus), and rats (both Rattus rattus (Linnaeus) and R. norvegicus (Berkenhout)). Feral pigs (Sus scrofa Linnaeus), which have also recently been demonstrated to eat large numbers of native lizards (J. Reardon pers. comm.), are widespread and abundant. Te Paki was one of the last areas of the country reached by possums, which have become abundant only in the last 20 years. In 1985 they were still recorded as absent but were spreading rapidly northward up the Aupōuri Peninsula, where they had reached about 10 km south of Waitiki Landing (Mitchell 1985). The relatively abundant (c, Published as part of Hitchmough, Rodney A., Nielsen, Stuart V., Lysaght, Judith A. & Bauer, Aaron M., 2021, A new species of Naultinus from the Te Paki area, northern New Zealand, pp. 389-400 in Zootaxa 4915 (3) on pages 391-398, DOI: 10.11646/zootaxa.4915.3.7, http://zenodo.org/record/4456586, {"references":["Robb, J. & Hitchmough, R. A. (1980) Review of the genus Naultinus Gray (Reptilia: Gekkonidae). Records of the Auckland Institute and Museum, 16, 189 - 200.","Hitchmough, R. A., Bull, L. & Cromarty, P. (Comps.) (2007) New Zealand threat classification system lists 2005. Department of Conservation, Wellington, 194 pp.","Bull, P. C. & Whitaker, A. H. (1975) The amphibians, reptiles, birds and mammals. In: Kuschel, G. (Ed.), Biogeography and Ecology in New Zealand. Junk, The Hague, pp. 231 - 276. https: // doi. org / 10.1007 / 978 - 94 - 010 - 1941 - 5 _ 5","van Winkel, D., Baling, M. & Hitchmough, R. (2018) Reptiles and Amphibians of New Zealand, a Field Guide. Auckland University Press, Auckland, 366 pp.","Nielsen, S. V., Bauer, A. M., Jackman, T. R., Hitchmough, R. A. & Daugherty, C. H. (2011) New Zealand geckos (Diplodactylidae): cryptic diversity in a post-Gondwanan lineage with trans-Tasman affinities. Molecular Phylogenetics and Evolution, 59, 1 - 22. https: // doi. org / 10.1016 / j. ympev. 2010.12.007","Nelson, N. J., Hitchmough, R. A., Monks, J. M. (2014) New Zealand reptiles and their conservation. In: Stow, A., Holwell, G. & Maclean, N. (Eds.), Austral Ark. Cambridge University Press, Cambridge, pp. 382 - 404. https: // doi. org / 10.1017 / CBO 9781139519960.020","Brook, F. J. & Thrasher, G. P. (1991) Cretaceous and Cenozoic geology of northernmost New Zealand. New Zealand Geological Survey Record, 41, 1 - 42.","Lux, J., Holland, W., Rate, S. & Beadel, S. (2009) Natural areas of Te Paki Ecological District: reconnaissance survey report for the Protected Natural Areas Programme. Department of Conservation, Northland Conservancy, Whangarei, viii + 319 pp.","Elliot, M. B., Striewski, B., Flenley, J. R. & Sutton, D. G. (1995) Palynological and sedimentological evidence for a radiocarbon chronology of environmental change and Polynesian deforestation from Lake Taumatawhana, Northland, New Zealand. Radiocarbon, 37, 899 - 916. https: // doi. org / 10.1017 / S 0033822200014983","Mitchell, N. (1985) The northward movement of possums. New Zealand Ecological Society Newsletter, 43, 11 - 12.","Dymock, J. J. (2000) Risk assessment for establishment of polistine (Polistes spp.) and vespine (Vespula spp.) wasps on the Three Kings Islands in the far north of New Zealand. In: Science for Conservation. Vol. 156. Department of Conservation, Wellington, pp. 1 - 18."]}