Kuniesaurus Albiauris, a New Genus and Species of Scincid Lizard From

Kuniesaurus albiauris, a New Genus and Species of Scincid Lizard from the Île des Pins, New Caledonia, with Comments on the Diversity and Aﬀinities of the Region’s Lizard Fauna Ross Sadlier, Matthias Deuss, Aaron Bauer, Hervé Jourdan

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Ross Sadlier, Matthias Deuss, Aaron Bauer, Hervé Jourdan. Kuniesaurus albiauris, a New Genus and Species of Scincid Lizard from the Île des Pins, New Caledonia, with Comments on the Diversity and Aﬀinities of the Region’s Lizard Fauna. Pacific Science, University of Hawaii Press, 2019,73(1), pp.123-141. 10.2984/73.1.6. hal-02496892

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HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Kuniesaurus albiauris, a New Genus and Species of Scincid Lizard from the Île des Pins, New Caledonia, with Comments on the Diversity and Affinities of the Region’s Lizard Fauna1

Ross A. Sadlier,2,6 Matthias Deuss,3 Aaron M. Bauer,4 and Hervé Jourdan5

Abstract: A new genus and species of skink, Kuniesaurus albiauris, is here described from the Île des Pins off southern New Caledonia. This new taxon possesses a unique suite of morphological apomorphies (scalation) that does not allow it to be placed in any existing Australasian eugongylid genus. It is known only from a single area in dense coastal forest on limestone, on the main island of the Île des Pins. The species area of occupancy is restricted, and the habitat occupied under threat from the spread of the highly invasive Little Red Fire Ant Wasmannia auropunctata. These factors place this new skink at a level of risk sufficient for it to be listed as Critically Endangered under IUCN Red List criteria. The affinities of the lizard fauna of the Île des Pins with respect to that present on southern New Caledonia are also discussed, most notably the absence from the Île des Pins of taxa typically restricted to ultramafic surfaces.

Keywords: Scincidae, systematics, new species, biogeography, conservation

THE ÎLE DES PINS is a moderately large island of small satellite islands. These islands are (∼150km2), 50km south of New Caledonia’s raised fossil coral heads formed ca. 118,000 B. main island, the Grande Terre, from which it P. ( Dubois et al. 1974) and mostly emergent is separated by a shallow series of straits, reefs, since the end of the Pleistocene (Balouet and lagoons, and passes. Although currently Olson 1989). They reach no more than 20 m separated, these two landmasses have experi- in elevation. enced repeated land-bridge connection and Field research on the Île des Pins in 1990 disjunction with fluctuating sea level over the by Bauer and Sadlier resulted in the first Pleistocene period (Pelletier 2007). Sur- comprehensive overview of the island’s lizard rounding the Île des Pins itself are a number fauna (Bauer and Sadlier 1994), but was both short in duration and limited in scope, being largely restricted to coastal habitats on the west side of the island. A comprehensive study 2Australian Museum Research Institute, Australian of the lizards of the satellite islands surround- Museum, Sydney, Australia. ing Île des Pins was undertaken by Geneva 3Independent naturalist—[email protected]. between 2000 and 2003, resulting in an 4Department of Biology, Villanova University, 800 updated overview of the region’s herpetofauna Lancaster Avenue, Villanova, Pennsylvania 19085, USA. (Geneva et al. 2013). Independent studies by 5Aix-Marseille Université, Université of d’Avignon, Ineich of the Muséum National d’Histoire CNRS, IRD, IMBE (Institut Méditerranéen de Biodiver- Naturelle, Paris (MNHN) resulted in the sité et d’Ecologie marine et continentale), Centre IRD Nouméa—BPA5, 98848 Nouméa Cedex, New Caledonia. rediscovery of the giant skink Phoboscincus 6Corresponding author (e-mail: ross.sadlier@ bocourti on one of the small islets (Ineich 2009) bigpond.com). after the species had been “lost” to science for nearly 150 years. These field studies in combination resulted in the recognition of a moderately rich lizard fauna of nine skinks and eight diplodactylid geckos (Geneva et al. 2013). More recently (2011–2014) field surveys by the Institut Méditerranéen de tails only, but estimated here as both speci- Biodiversité et d’Ecologie marine et con- mens had regrown tails. Body measurements tinentale (IMBE) lab team (HJ coordinator), (axilla to groin, forelimb to snout, hindlimb, supported by the Province Sud, have been and tail lengths) are for both specimens, and undertaken to document the extent and are expressed as a percentage of snout-to-vent impact of invasive species (rodents, feral cats, length in the taxon account. Sex of the and exotic ants) on the satellite islands holotype as female was determined by the surrounding Île des Pins, and to a lesser presence of eggs, and the paratype as male by extent the main island, with an emphasis on the presence of testes. their impact on the region’s lizard fauna. Scalation. Midbody scale rows (MBR)— These field studies have resulted in significant number of longitudinal scale rows around new records for the offshore islands in the body counted midway between axilla and region, and also investigated forest remnants groin; dorsal scale rows (DSR)—number of on ultramafic surfaces of the main island scales in a row from first scale posterior to (Jourdan et al. 2011, 2012, 2013, and 2014). parietal scale to last scale at level of vent In 2015, a small skink, clearly representing opening; fourth finger (FFS) and toe (FTS) an undescribed taxon, was found opportunis- scales—number of dorsal scales on fourth tically on the main island, in the Touete tribal digit of the manus and pes, distal scale region. Most significantly, this new species contains claw, basal scale broadly contacts could not be assigned to any of the existing adjacent basal scale of third finger or toe; genera of New Caledonian scincid lizards, or fourth finger (FFL) and toe (FTL) lamellae— to any existing Australasian genus. The number of ventral scales on fourth digit of the discovery of this uniquely divergent species, manus and pes, distal scale contains claw, basal indicating that the evolutionary history of the scale is last largely undivided scale at, or lizard fauna of the Île des Pins is far more proximal to, a point level with intersection of complex than initially thought, contributes third and fourth digits. Bilateral scalation significantly to the region’s biodiversity value. characters were scored on both sides and the The suite of morphological apomorphies in mean value used. scalation unique to the new species identify it The description of the new species pre- as an evolutionary lineage distinct from other sented here is based on two specimens, the genera of skinks, and a new genus is required holotype and paratype, with variation in to accommodate it. coloration supplemented by images of live individuals not vouchered. Allocation of the new species to a new genus is based on MATERIALS AND METHODS morphological criteria, with comparisons Abbreviations: Institutional abbreviations— made to other genera in the Eugongylus group MNHN (Muséum National d’Histoire Nat- of skinks, a morphologically based lineage as urelle, Paris); IMBE (Institut Méditerranéen defined by Greer (1979) for which monophyly de Biodiversité et d’Ecologie marine et has since been independently corroborated in continentale). subsequent genetic studies (Honda et al. Morphological Comparisons: Measure- 2000). Tissue samples were not available to ments. Snout to vent length (SVL)—measured generate DNA sequence data for inclusion of from tip of snout to caudal edge of anal scales; the new species into the framework of axilla to groin distance—measured from phylogenetic relationships for New Caledo- middle of base of forelimb to middle of base nian skinks derived from molecular data of hindlimb; forelimb to snout length— (Smith et al. 2007). This study identified measured from tip of snout to middle of base the endemic New Caledonian skink genera as of forelimb; hindlimb length—from middle of part of a regionally discreet monophyletic base of hindlimb to tip of fourth toe including evolutionary lineage, the Tasmantis group of nail; tail length—measured from caudal edge skinks, one that also includes the New of anal scales to tip of tail on complete original Zealand skinks and the endemic species found on Lord Howe Island and the Norfolk Island Kuniesaurus gen. nov. Sadlier, Deuss, (s) group in the southwest Pacific Ocean. and Bauer Monophyly of the Tasmantis group was Type species: Kuniesaurus albiauris n. sp. supported in a later genetic study, by Chapple Diagnosis: Kuniesaurus can be distin- et al. (2009), which focused on the New guished from all other genera in the Eugon- Zealand skink fauna. However, the study by gylus group of Greer (1979) by the following Smith et al. (2007) and a later analyses of combination of derived character states molecular data for the New Caledonian skinks (apomorphies) as defined by Sadlier (2010): by Ineich et al. (2014) have both supported the supranasal scale or postnasal suture absent; robustness of genera proposed primarily on anterior loreal reduced in size and present as a morphological criteria to accommodate semilunar scale positioned on the poster- the diversity of evolutionary lineages within odorsal margin of the nasal and failing to the endemic New Caledonia skink fauna. The contact the upper labials; subocular scale row only exception to this was Lioscincus, which complete; a reduction in the number of upper was resurrected from synonymy by Bauer and labial scales (to six) and lower labial scales (to Sadlier (1993) to accommodate a suite of five); frontoparietal scales fused; lower eyelid residual New Caledonian taxa formerly with an obvious, centrally located semitran- included in Leiolopisma and explicitly identi- sparent disc; ear lobules barely distinguishable fied as polyphyletic (Sadlier et al. 1998), a from blunt conical scales around upper, lower, concept later supported in the molecular and posterior edges of ear opening; diminu- phylogeny presented by Smith et al. (2007). tive size (adults <50mm SVL). Subsequently, Sadlier et al. (2015) used a The New Caledonian taxa Celatiscincus and combination of morphological and genetic Marmorosphax share the greatest number of information to delineate four monophyletic derived character states with Kuniesaurus, genera from among its constituent species. namely the lack of a supranasal scale or The only member of the New Caledonian postnasal suture; anterior loreal significantly lizard fauna for which significant phylogenetic reduced in size (failing to contact the upper uncertainty remains is Geoscincus haraldmeieri, labials in Marmorosphax); in having a complete an unusual species known only from the two subocular row of scales; frontoparietal scales individuals included in the original descrip- fused; lower eyelid with an obvious, centrally tion in 1976 (Böhme 1976; Sadlier 1986). located semitransparent disc; and a large round ear opening with the lobules barely distinguish- SYSTEMATICS able from adjacent scales. Kuniesaurus has a The following suite of morphological char- reduced number of upper labialand lower labial ∗ acteristics (apomorphies ) identify the new scales, derived character states not present in species of lizard described here from the Île either Celatiscincus or Marmorosphax, both of des Pins as a member of the Eugongylus group which have the modal plesiomorphic state of skinks as defined by Greer (1979): parietal (upper labials seven; lower labials six). Both ∗ scales meeting behind the interparietal ; Celatiscincus and Marmorosphax have keeled parietal bordered along its posterior edge by body scales, a trait considered apomorphic an upper secondary temporal and transversely within the New Caledonian skink radiation, ∗ enlarged nuchal scale ; medial preanal scales whereas Kuniesaurus has smooth body scales, more or less subequal in size to and over- the state considered plesiomorphic within the lapped by more lateral preanals; and the scales New Caledonian skink radiation. Celatiscincus on the dorsal surface of the fourth toe are in a and Marmorosphax also each possess additional single row throughout the length of the digit. apomorphies not shared with Kuniesaurus.In The number of premaxiliary teeth and other Celatiscincus the parietal scales are each bor- aspects of osteology have not been assessed on dered by a single enlarged temporal scale and the limited number of specimens from which divided nuchal scale into two (sometimes three) the species is known. scalesthatare transversely enlargedbutequalin FIGURE 1. Adult female holotype of Kuniesaurus albiauris n. sp. (MNHN-RA-2017.0101) showing the darkly streaked dorsal and lateral color pattern of the body (A) and side of head (B). size to the adjacent body scales (versus the Recognized species: Kuniesaurus albiauris n. plesiomorphic condition of the parietals each sp. bordered by a single enlarged temporal scale and transversely enlarged nuchal scale in Sadlier, Deuss, Bauer, Kuniesaurus), and in Marmorosphax the division Kuniesaurus albiauris of the third pair of chinshields such that the and Jourdan n. sp. scales contacting the lower labials are separated Isle of Pines white eared skink/Scinque à from each other by an intervening row of five oreilles blanches de l’Île des Pins. scales (versus the plesiomorphic condition of Figures 1–5 the enlarged third pair of chinshields separated Type Material: Holotype MNHN-RA- from each other by three intervening scales in 2017.0101 New Caledonia, Île des Pins Kuniesaurus). 22.59748° S 167.52472° E(Figure 1 and Kuniesaurus has an egg-laying mode of Figure 2). Paratype MNHN-RA-2017.0102 reproduction (oviparous the plesiomorphic New Caledonia, Île des Pins 22.59427° S condition), a trait that further distinguishes it 167.52453° E—both holotype and paratype from Marmorosphax, which is live-bearing collected by Matthias Deuss, 20 December (ovoviviparous the apomorphic condition), at 2015. least for those species in which the mode of Etymology: The species epithet albiauris is reproduction has been determined. a noun in apposition and is derived from the Etymology: The name is derived from Latin albus (white) and auris (ear) and is used Kunié, given to the main island of the Île des in reference to the prominent white spot in Pins by its indigenous Kanak population, and the region of the lower secondary temporal saurus, the New Latin form of the Greek giving the appearance of having white ears. sauros=lizard. The gender is masculine. Diagnosis: see generic diagnosis. FIGURE 2. Head scalation of adult female holotype of Kuniesaurus albiauris n. sp. (MNHN-RA-2017.0101), showing the lateral (upper), dorsal (middle), and ventral (lower) aspects. FIGURE 3. Adult male paratype of Kuniesaurus albiauris n. sp. in life.

FIGURE 4. Dorsal (A) and ventral (B) surface of adult female holotype (MNHN-RA-2017.0101 –- right) and adult male paratype (MNHN-RA-2017.0102 – left) Kuniesaurus albiauris n. sp. prior to preservation, note the undersurface of the head and neck are pale, graey in color, whereas the undersurface of body and tail (B) has a yellow flush posterior of the forelimb. FIGURE 5. Adult Kuniesaurus albiauris n. sp. (A) (sex unknown) from the type locality, note the distinctive white temporal spot on the side of the head, and a juvenile (B) (sex unknown) from the same area. Description: Scalation and coloration is major head shields and a concentration of dark based on the single adult female holotype and markings on the supraciliary shields. Lateral adult male paratype. surface of the body with an underlying Measurements. Snout to vent length (SVL) background color that grades from brown 46.0mm (holotype) 40.5mm (paratype); dis- uppermost (an extension of the dorsal color) tance from axilla to groin 54.3% of SVL; through to gray approaching the ventrolateral distance from forelimb to snout 39.5%–40.2% surface, extensively marked with a pattern of ofSVL;hindlimblength29.3%–33.3%ofSVL; dark longitudinal streaks (similar to the dorsal tail lost in both individuals but from images of surface) and to a lesser extent scattered pale live specimens would appear to be approxi- spots. Width of the dark streaks variably mately equal in length to body ∼100% SVL. dominates each scale, but is particularly Scalation. Prefrontals large, narrowly to intense on the upper lateral surface where it moderately separated; frontoparietals fused; approaches the dorsal surface, resulting in interparietal distinct; parietals meet behind most scales in the dorsolateral region having a interparietal, parietals each bordered by a dark median streak. Paler areas of of the transversely elongate upper secondary tem- posterior half of the lateral surface of the body poral scale and nuchal; primary temporal with a dull yellow flush overall in life (similar single; upper secondary temporal single; lower to adjoining ventral surface), including the secondary temporal single; tertiary temporals pale spots of the upper lateral surface of the two; postlabials scales two; nasal scales mod- body, neck and head, whereas the pale mid and erately large, moderately/widely separated; lower lateral areas of the anterior part of the supraciliaries seven; upper labials six with the body either side of the forelimb are gray to fourth subocular and separated from contact pinkish-gray in life. Side of head with an with the lower eyelid by a complete row of underlying background color of light brown subocular scales; lower labials five, first two to gray similar to the lateral surface of the contacting postmental; enlarged chinshields body, but dark overall by virtue of the head three, members of first pair in broad contact, scales being dominated by extensive dark members of second pair separated by one scale, markings, and with a pattern of pale spots and members of third pair by three scales. Lower blotches to the labial and temporal scales, eyelid with an obvious, centrally located, these with a faint yellow flush in life except for semitransparent disc. Ear opening large and the large and prominent spot on the lower without obviously enlarged lobules anteriorly. secondary temporal scale, which is white. The Body scales smooth. Midbody scale rows 32; undersurface of the head and neck is pale gray dorsal scale rows 53–56; scales on top of fourth in life, and with large dark blotches present on finger 6–8(x ¼ 7); lamellae beneath fourth the outer margin of the postmental scale and finger 12–14 (x ¼ 11:6); scales on top of fourth the chinshields, where they contact the toe 11; transversely enlarged lamellae beneath adjoining dark blotches of the lower labials. fourth toe 20–23 (x ¼ 21:25), with a series of Undersurface of body and tail is pale, in life several granular scales basally. with a yellow flush posterior of the forelimb, Coloration. Adult female (Figure 1 and and the tail has a row of dark spots on the Figure 4A right and 4B right): Dorsal surface underside along the ventrolateral margin. of the body brown with a series of dark broken Adult male (Figure 3 and Figure 4A left and longitudinal stripes down the body from the 4B left): Dorsal surface of the body brown nape to past the level of the hindlimbs. Each with occasional scattered dark markings down stripe composed of a series of prominent dark the center of the dorsal scales. Lateral surface streaks, each of which runs down the center of of the body with an underlying background the dorsal scale (often failing to reach the color of cream to gray with a pattern of anterior and/or posterior edge of the scale) irregularly distributed dark longitudinal and can be absent from each second or third streaks uppermost, becoming more diffuse scale. Dorsal surface of head mid-brown over the remainder of the side of the body, but (similar to body) with dark blotches on most also with scattered dark blotches. Dorsal surface of head brown (similar to body) with region of the lower secondary temporal only a few dark markings, and the side of head (Figure 5A), and a dull golden yellow color with an underlying background color of gray on the posterior part of the rim of the lower but with each scale dominated by extensive eyelid, and to a lesser extent the posterior part dark markings, and the same pattern of pale of the upper eyelid. spots and blotches to the labial and temporal Juveniles (Figure 5B): The only juvenile scales as in the adult female, dull cream to specimen photographed (non-vouchered and faint yellow in life except for the large and of undetermined sex) was very dark, with very prominent pale spot near the lower secondary faint yellow streaks on jaws, almost no color temporal scale, which is white. The under- transition between dorsum and sides, more surface of body, neck, and head similar to the conspicuous dark markings on ventral side, adult female (above), also with a yellow flush including between jaws, forelimbs and hin- on the abdomen posterior of the forelimb and dlimbs, and the pale yellow area on ventral side along the underside of the tail (Figure 4B left). extended frontward further than in adults. Images of other live adult individuals from Distribution and Habitat: Very little is Île des Pins (sex unknown) also show these to known of the natural history of this species. have a yellow flush to the abdomen posterior The only known area of occurrence is in humid of the forelimb and along the underside of the forest on calcareous soils in the northeastern tail, a large and prominent white spot in the part of mainland Île des Pins (Figure 6). Here

FIGURE 6. Location of the type locality for Kuniesaurus albiauris n. sp. (area within dashed lined) on the Île des Pins. Bright green areas represent forests on coralline soil, whereas dark green areas represent forests on ultramafic soils. Source of data: Gouvernement de la Nouvelle-Calédonie. Figured by Matthias Deuss. FIGURE 7. Habitat from which Kuniesaurus albiauris n. sp. was collected in humid forest on coralline soils on the Île des Pins. the species was frequently encountered shel- tions are more humid. It has not been recorded tering under rocks and decaying wood along a from the adjacent, near sea level, disturbed track (Figure 7), and when uncovered quickly littoral forest, which is dominated by Cocos escaped beneath the litter; it has not been nucifera and Araucaria columnaris. Kuniesaurus observed active on the surface. This pattern of has been encountered in three different behavior is similar to the species of Marmoro- periods: June, October, and December during sphax which are absent from Île des Pins, and cloudy to sunny and hot weather, whereas Kuniesaurus albiauris n. sp. may fill a similar sympatric Sigaloseps sp. was not found in June ecological niche. At this location it is sympatric in a three-hour search. with five other small species of skink. Of these, Reproductive biology: This species is Nannoscincus sp. are almost exclusively fossorial oviparous, the adult female holotype collected in habits and Sigaloseps sp. primarily fossorial in December contained three large shelled but with periods of surface activity, Celatiscincus eggs. One juvenile was observed in June 2016. euryotis appears to be a litter dwelling species Conservation status: Kuniesaurus albiauris with frequent periods of surface activity (HJ n. sp. has a very restricted known area of and MD pers. obs.), and the species Caledo- occupancy. What is known of its distribution niscincus bodoi and Caledoniscincus atropunctatus on the island indicates it occupies the less are primarily surface active litter skinks and disturbed forest on uplifted coralline surfaces both are common at the type locality. Based on on the east side of the island. The area of the few surveys carried out on this track, humid forest on calcareous soils on the Île des Kuniesaurus seems restricted to the uplifted Pins is estimated at around 75 km2, approxi- coralline “plateau” (like Celatiscincus euryotis) mately half the surface of the island (not where the forest is less disturbed and condi- including Ile Kotomo), although the actual extent of suitable forest habitat is likely to be unreported high level of cat predation on less, given some areas of forest on the west native skinks, especially the ground dwelling side of the island are likely to be disturbed by species Marmorosphax tricolor. human activities and represent areas of The species restricted extent of occur- regrowth forest/secondary forest. rence, in combination with the threats posed The most significant threats to Kuniesaurus by the presence of rodents and the occurrence albiauris n. sp. come from the presence of of the invasive ant Wasmannia auropunctata invasive species, primarily rodents (Pacific Rat near its only known site, place Kuniesaurus and Ship Rat), feral cats, and invasive ants. albiauris n. sp. at high level of risk, satisfying The negative impact of rats on island lizard the criteria to be ranked as Critically populations has been well documented (Whi- Endangered under the IUCN Red List taker 1973; Monks et al. 2014), with their classification system (IUCN 2017): it has a presence clearly suppressing potential lizard its extent of occurrence is estimated to be less population densities (Thibault et al. 2017). A than 100 km2 (criteria B1); it is known only negative impact on lizard populations by the from one location (criteria B1a); and by virtue Little Red Fire Ant (LFA) Wasmannia aur- of the impact of introduced predators it has an opunctata has also been identified, with studies inferred decline in area, occurrence, occu- in dry forest habitat in New Caledonia pancy and extent and/or quality of habitat reporting a lower abundance of several lizard (criteria B1b i–iii). The population status of species in invaded areas (Jourdan et al. 2001), the species at the type locality needs to be and it is considered to be a major threat island- investigated, as does its possible occurrence in wide. Similarly, the presence of the introduced suitable habitat elsewhere on the Île des Pins Yellow Crazy Ant (YCA) Anoplolepis gracilipes and its surrounding islands. could also present a threat. Booms in Crazy Ant populations have been identified as a likely contributing factor in the decline of DISCUSSION some lizard species on Christmas Island in the The rediscovery of the giant skink Phoboscincus Indian Ocean (Smith et al. 2012). Both the bocourti in 2003 (Ineich 2009) on islets LFA and YCA are reported from coastal surrounding the Île des Pins, almost 150 years limestone forests on Île des Pins. At this time after it was initially discovered, is one among the YCA is more established in the forest on the following factors, all indicating that the Île the west coast of the main island, whereas LFA des Pins and its associated islets have a richer is actively spreading on the east coast, and has and more complex lizard fauna than previously been detected in close proximity to the area of thought: the formal recognition of the giant the known population of Kuniesaurus (Jourdan gecko Rhacodactylus trachycephalus as an island et al. 2011, 2012, 2013). The spread of the endemic (Bauer, Jackman et al. 2012), the LFA is a major concern for long-term survival recognition of the skinks Celatiscincus euryotis of Kuniesaurus given it is a cryptic species that (Sadlier, Smith et al. 2006) and Caledoniscincus inhabits litter, where the LFA is known to bodoi (Sadlier, et al. in prep) as regional island establish nests. The threats posed by these endemics, and now the discovery of a mono- ants come from the physical displacement of typic genus of skink endemic to the region and lizards from sheltering and foraging sites by two other species of skink not previously the presence of large numbers of ants during recorded (an undetermined species of Sigaloseps boom periods, and indirectly from the loss of and Nannoscincus). potential invertebrate prey that constitute Of particular interest is the composition of part of the lizards diet, as a result of declines the lizard fauna present on the Île des Pins and accompanying the spread of the invasive ants its satellite islands with respect to that of the (Jourdan et al. 2001). The negative impact of southern Grande Terre, the region south of feral cats on the lizard fauna of New the Thio Valley dominated by the southern Caledonia has recently been recognized. ultramafic bioregion (Figure 8), as well as the Palmas et al. (2017) illustrated a previously inference that can be drawn regarding their FIGURE 8. The area of ultramafic surfaces (light green) covered by the southern ultramafic bioregion in southern New Caledonia, showing the northern limits at the Thio valley (dash-dot-dash brown line) and the area in the south referred to as the Grand Sud, a sub-region within which a number of southern ultramafic endemic species appear to be restricted. Source of data: DIMENC (direction de l’industrie, des mines et de l’énergie), Gouvernement de la Nouvelle- Calédonie, 2012. historical distribution and the nature of past scenario is seen in the insular endemic connectivity between the two areas. Celatiscincus euryotis and its only congener, The occurrence on both the Île des Pins C. similis from the northern Grande Terre and in southern New Caledonia of lizard (Sadlier, Smith et al. 2006). The morpholo- species that lack obvious water-crossing gical distinctiveness of Kuniesaurus albiauris n. capabilities, such as the geckos Correlophus sp., with no apparent obvious close relation- ciliatus, Mniarogekko chahoua, and Eurydacty- ship to any known genus, hints at an even lodes vieillardi, suggests that land connection deeper history of connectivity and isolation sufficient for dispersal of such lizards existed between the Île des Pins and southern New historically between the two regions, and the Caledonia. presence of two lineages each of which has The southern Grande Terre has a rich highly divergent sister taxa consisting of a lizard fauna of 43 species endemic to New taxon endemic to the Île des Pins with an Caledonia. Extensive field research in this evolutionary counterpart on Grande Terre, region over the past 10 to 15 years has greatly further suggests that historically older pat- increased our knowledge of the distribution terns of connection and isolation between the and habitat preferences of many species in this two regions also existed prior to those of the region, and resulted in the discovery of a Pleistocene period identified by Pelletier number of new taxa (Bauer et al. 2008; Bauer, (2007). The high level of genetic differentia- Sadlier et al. 2012; Sadlier, Bauer et al. 2006; tion between the regional endemic Rhacodac- Sadlier et al. 2013, 2014a, 2014b, 2014c). Over tylus trachycephalus and its sister species R. half of the lizard species recorded are regional trachyrhynchus (Bauer, Jackman et al. 2012) endemics and most are restricted to habitats indicates that they have long been separated on ultramafic surfaces, identifying it as a from one another. A similar evolutionary significant bioregion for lizards, with the remainder tending to be more widespread in and maquis shrubland) and nonultramafic distribution and also occurring on habitats on surfaces (robusta coastal forest, sclerophyll non-ultramafic surfaces (Ta b l e 1 ). Of the 17 forest and mangroves; sauvagii sclerophyll native species now recorded from the Île des forest). Pins (excluding the as yet undetermined species Six of the diplodactylid geckos recorded of Sigaloseps and Nannoscincus), 12 also occur on from southern New Caledonia have not been the southern Grande Terre (some more widely recorded on the Île des Pins: five of these are distributed); the remaining 5, Rhacodactylus endemic to the southern ultramafic bioregion, trachycephalus, Caledoniscincus bodoi, Celatiscincus and one, Rhacodactylus auriculatus, is restricted euryotis, Kuniesaurus albiauris n. sp., and to ultramafic surfaces of both southern and Phoboscincus bocourti, are endemic to the island northern Grande Terre. Twoof these (Bavayia region (Ta b l e 1 ). Given the increase in knowl- geitiana and Eurydactylodes symmetricus)are edge of the lizard fauna of both the southern widespread in the bioregion, one (Correlophus Grande Terre and the Île des Pins, we take the sarasinorum) is widespread in the south of the opportunity to review the affinities of the bioregion, while the remaining two have more species found in both regions. restricted distributions, one (Bavayia nubila)to Seven species of diplodactylid geckos are the interior ranges of the southern ultramafic recorded as native to the Île des Pins (Geneva region and one (Bavayia goroensis) to the ultra- et al. 2013), including four species of giant mafic plateau in the far south of the bioregion. gecko (Correlophus ciliatus, Mniarogekko cha- While the two range-restricted species likely houa, Rhacodactylus leachianus, and Rhacodacty- represent localized endemics, the absence of lus trachycephalus), two species of Bavayia the other four species from the Île des Pins is (Bavayia robusta and Bavayia sauvagii), and less readily explained, given their presence in one Chameleon Gecko (Eurydactylodes vieil- ultramafic habitats on the adjacent southern lardi). All except one, the giant gecko Grande Terre. Rhacodactylus trachycephalus, also occur on The composition of the skink fauna of Grande Terre (Table 1). Three of the giant southern Grande Terre relative tothatrecorded geckos recorded from the main island of the from the Île des Pins is even more complex. An Île des Pins, Correlophus ciliatus, Mniarogekko extensive suite comprising 28 species of skink is chahoua, and Rhacodactylus leachianus, also known to occur in the southern Grande Terre. have broad (but patchy in ciliatus and chahoua) Some species, as currently conceived, are taxa distributions on the Grande Terre in mostly with widespread distributions on the Grande forested habitats on both ultramafic and Te r r e ( n=12), but the majority (16) are taxa nonultramafic surfaces. All three have been endemic to the southern ultramafic bioregion. recorded in the southern Grande Terre, Some of the skink taxa endemic to the southern primarily in humid forest on the southern ultramafic region have restricted distributions, ultramafic ranges, but Rhacodactylus leachianus either in the ranges of the interior (8) and/or the also occurs in other forest habitats, including far south—in the area known as the Grand Sud coastal forests and islets off the east coast. The (5)— but some are also widespread (4). With three other gecko species, Bavayia robusta, two possible exceptions (discussed below), none Bavayia sauvagii, and Eurydactylodes vieillardi, of the skink taxa endemic to the southern are also found on both the Île des Pins and the ultramafic bioregion are found on the Île des Grande Terre. The complexities of the Pins. systematics of both Bavayia robusta and Aside from the five island endemics (includ- Bavayia sauvagii are still under review but ing Kuniesaurus albiauris n. sp.) the skinks the taxa ultimately to be recognized under recorded from the Île des Pins also includes six these names will be restricted to southern taxa that are widely distributed on Grande Grande Terre including the Île des Pins. Terre. Twoadditional species, one in the genus Across their range they occupy a range of Sigaloseps and one in the genus Nannoscincus,are habitats on both ultramafic (robusta humid also known to occur on the Île des Pins, but the forest; sauvagii humid forest, maquis forest identity of these and whether they are con- TABLE 1 Skinks and geckos recorded from southern Grande Terreand Île des Pins showing those with distributions that extend across the Grande Terre (GT), those with distributions limited to the southern Grande Terre but recorded from both ultramafic and non-ultramafic surfaces (SGT); those restricted to ultramafic surfaces of the southern ultramafic bioregion (SUB); and taxa recorded from the Île des Pins (IoP). A general assessment of the species distribution is also given.

Diplodactylidae Bavayia geitaina SUB Widespread across SUB Diplodactylidae Bavayia goroensis SUB Restricted to south of SUB Diplodactylidae Bavayia nubila SUB Restricted to ranges of SUB Diplodactylidae Bavayia robusta SGT IoP Widespread across SGT+IoP Diplodactylidae Bavayia sauvagii SGT IoP Widespread across SGT+IoP Diplodactylidae Bavayia septuiclavis SUB Widespread across SUB Diplodactylidae Correlophus ciliatus GT IoP Widespread across GT+IoP Diplodactylidae Correlophus sarasinorum SUB Restricted to south of SUB Diplodactylidae Eurydactylodes symmetricus SUB Widespread across SUB Diplodactylidae Eurydactylodes vieillardi GT IoP Widespread across GT+IoP Diplodactylidae Rhacodactylus auriculatus GT Widespread across GT Diplodactylidae Rhacodactylus trachycephalus IoP Restricted to IoP Diplodactylidae Rhacodactylus trachyrhynchus GT Widespread across GT Diplodactylidae Rhacodactylus leachianus GT IoP Widespread across GT+IoP Diplodactylidae Mniarogekko chahoua GT IoP Widespread across GT+IoP? Scincidae Caesoris novaecaledoniae GT Widespread across GT Scincidae Caledoniscincus atropunctatus GT IoP Widespread across GT+IoP Scincidae Caledoniscincus austrocaledonicus GT Widespread across GT Scincidae Caledoniscincus bodoi IoP Restricted to IoP Scincidae Caledoniscincus festivus GT Widespread across GT Scincidae Caledoniscincus haplorhinus GT IoP Widespread across GT+IoP Scincidae Caledoniscincus notialis SUB Widespread across SUB Scincidae Celatiscincus euryotis IoP Restricted to IoP Scincidae Cryptoblepharus novocaledonicus GT IoP Widespread across GT+IoP Scincidae Epibator greeri GT Widespread ? across GT Scincidae Epibator nigrofasciolatus GT IoP Widespread across GT+IoP Scincidae Graciliscincus shonae SUB Restricted to south of SUB Scincidae Kuniesaurus albiauris n. sp. IoP Restricted to IoP Scincidae Lacertoides pardalis SUB Widespread ? across SUB Scincidae Marmorosphax montana SUB Restricted to ranges of SUB Scincidae Marmorosphax tricolor GT Widespread across GT Scincidae Nannoscincus fuscus SUB Restricted to ranges of SUB Scincidae Nannoscincus garrulus SUB Restricted to ranges of SUB Scincidae Nannoscincus gracilis GT Widespread across GT Scincidae Nannoscincus mariei SUB Restricted to south of SUB Scincidae Phaeoscincus ouinensis SUB Restricted to ranges of SUB Scincidae Phasmasaurus tillieri SUB Widespread across SUB Scincidae Phoboscincus bocourti IoP Restricted to IoP Scincidae Phoboscincus garnieri GT IoP Widespread across GT+IoP Scincidae Sigaloseps deplanchei SUB Restricted to south of SUB Scincidae Sigaloseps ruficauda SUB Restricted to ranges of SUB Scincidae Sigaloseps balios SUB Restricted to ranges of SUB Scincidae Sigaloseps ferrugicauda SUB Restricted to ranges of SUB Scincidae Sigaloseps pisinnus SUB Restricted to ranges of SUB Scincidae Sigaloseps conditus SUB Restricted to ranges of SUB Scincidae Simiscincus aurantiacus SUB Restricted to south of SUB Scincidae Tropidoscincus aubrianus GT IoP Widespread across GT+IoP Scincidae Tropidoscincus variabilis SUB Widespread across SUB

Note: The presence of Epibator greeri in southern New Caledonia is based on photographic images (Deuss, unpublished; VALE Nouvelle-Calédonie unpublished; Jean-Louis Ruiz), and similarly that of Caesoris novaecaledoniae is based on an image on the Endemia website of an individual from Mont Dore (http://endemia.nc/faune). Note: The record of Bavayia crassicollis reported by Geneva et al. (2013) from Île des Pins is not included here as it is considered to likely represent a recent human-mediated introduction and not native to the region. specificwithtaxaknownfromGrandeTerrehas plateau and adjacent interior ranges in the far yet to be determined. Of the six species shared southofthe bioregion. The other two species of between southern Grande Terre and the Île des Nannoscincus have more localized distributions, Pins, one, Cryptoblepharus novocaledonicus,isa N. garrulus having only been recorded from coastal specialist (not a member of the high elevation forest habitat on ultramafic Tasmantis radiation, sensu Smith et al. 2007) surfaces in the far north of the bioregion, and four, Caledoniscincus haplorhinus, C. atro- whereas the population of N. gracilis in the punctatus, Epibator nigrofasciolatus,andPhobos- bioregion is known only from several low cincus garnieri,are“generalist” species with elevation sites on coastal non-ultramafic sur- widespread distributions in the territory and faces and the base of the ultramafic surfaces known to occur in a range of habitats (including near Nouméa and on the southwest coast. secondary associations) on both ultramafic and Aside from the two possible exceptions just non-ultramafic surfaces. The presence of these discussed, the number of species recorded “generalist” species on both the Île des Pins and from the south of the Grande Terre that are the Grande Terre is not surprising given these restricted to ultramafic surfaces, but absent taxa also occur on the Loyalty Islands, which from the Île des Pins, is extensive, and even have never shared a land connection to the when narrow-range localized endemic taxa Grande Terre, indicating they have good over- are excluded comprises nine skinks and four water dispersal abilities (although it is possible geckos, a number of which are found across a some might have been inadvertent introduc- broad range of habitat types on ultramafic tions). The remaining species found on the Île surfaces. One possible explanation to account des Pins that is also widespread on Grande for this anomaly is the absence of suitable Terre is Tropidoscincus aubrianus, an enigmatic habitat on the Île des Pins for many of the taxa species in that it appears to be a generalist in restricted to habitats on ultramafic surfaces. habitat preferences but has only been recorded The giant gecko Correlophus sarasinorum is sporadically, and has not been found on widespread in the south of the bioregion but ultramafic surfaces. The affinities of the absent from the Île des Pins. In southern Sigaloseps recently recorded on Île des Pins Grande Terre it has only been recorded from has yet to be determined. It would appear to be humid forest, and its absence from the Île des aligned with Sigaloseps deplanchei, a species Pins may lie in its preference for a particular generally regarded as restricted to habitats on type or level of development of humid forest the ultramafic surfaces of southern Grande not found there. The giant gecko Correlophus Terre (Sadlier et al. 2014a), but which has also ciliatus has been recorded from humid forest been recorded from coastal forest habitat habitat on the Île des Pins and in southern fringing ultramafic surfaces (Port Boise and Grande Terre, and seasonally dry vallicole Yate), these occurrences being regarded as a forest in the north of Grande Terre.Similarly, localized extension of populations resident on Mniarogekko chahoua is known from humid the nearby ultramafic surfaces. However, the forest habitat in southern Grande Terre and presence of a species of Sigaloseps on the Île des assumed to occur in this habitat type on the Île Pins in habitat on non-ultramafic surfaces des Pins; its counterpart in northern Grande indicatesthatthehabitatpreferencesforspecies Terre, Mniarogekko jalu, is also known to in the genus may be broader than initially inhabit seasonally dry vallicole forest. As such, thought. The implications for a sight record for the species C. ciliatus and M. chahoua would Nannoscincus on the Île des Pins (Deuss, pers. appear to be able to occupy a broader range of obs.) remain uncertain until the identity of the forest types, whereas Correlophus sarasinorum species can be determined. Four species of appears to be constrained to well-developed Nannoscincus occur in the southern ultramafic humid forest (Sadlier pers. obs.), subtleties in bioregion. Twoof these, N. fuscus and N. mariei, biology that may have contributed to these are known only from ultramafic surfaces, the species’ presence or absence on Île des Pins. former primarily from habitat on the ranges Although humid forest habitat is present and the latter from habitat on the ultramafic today on Île des Pins, the forest type that existed on the island and/or areas of emergent the apparent limitations associated with “land-bridges” connecting southern Grande evolution on ultramafic surfaces. However, Terre to the Île des Pins during historically these comments are made with the caveat that drier periods when sea levels were lower, may only limited survey work for lizards has been have been structurally different, and while (by undertaken on the island’s ultramafic surfaces, inference) able to provide the microhabitat and the only area that has been investigated in (sheltering and foraging sites) required by C. detail, Pic N’Ga, has undergone significant ciliatus and M. chahoua, may not have reached modification through a high level of defor- the level of development seen in humid forest estation and burning, with almost half the of southern Grande Terre as required by plateau area converted to pine plantation. Correlophus sarasinorum. Clearly, targeted and systematic survey work However, the absence from the Île des Pins for lizards is required on both the forest areas of a number of species restricted to ultramafic in the west of the island and on the remaining surfaces but with distributions that extend natural habitat on its ultramafic surfaces in across the whole of the southern ultramafic order to fully assess the diversity of the lizard region and occur in a range of both forest and fauna of the Île des Pins in a regional context, maquis habitats (Bavayia septuiclavis, Eurydac- and its role as a hotspot for lizards within the tylodes symmetricus, Caledoniscincus notialis, New Caledonian biodiversity hotspot. Tropidoscincus variabilis) may not only lie in the unsuitability of habitat present on the Île ACKNOWLEDGMENTS des Pins now or historically, or of that on historical land bridges that once connected The authors thank the Direction de l’Envir- the regions, but also in the response of lizards onnement Province Sud for the financial and endemic to southern ultramafic regions to logistic support provided to IRD to undertake drier conditions associated with historical research on the Île des Pins which included glacial periods. Contraction of range to mesic studies on the region’s lizard fauna, in refuges during historically drier periods has particular Isabelle Jurquet, Manina Tehei, been proposed as a process that has driven intraspecific lineage diversification of lizards Caroline Groseil and Cendrien Meresse. in the Wet Tropics region of Australia during AMB was supported by National Science the Pleistocene, and deeper species-level Foundation grant DEB 1555968 and by the diversification earlier in the Miocene (Schnei- Gerald M. Lemole Endowed Chair Funds der et al. 1998; Schneider and Moritz 1999; from Villanova University. Moritz et al. 2012). Similar processes have been proposed in the evolution of elements of Literature Cited the lizard fauna on the ultramafic surfaces of New Caledonia, leading to regionally and Balouet, J., and S. Olson. 1989. 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