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A new species of Underwoodisaurus (Squamata: Gekkota: Carphodactylidae) from the Pilbara region of Western Australia
Abstract and Figures
Ongoing surveys and systematic work focused on the Pilbara region in Western Australia have revealed the existence of numerous unrecognized species of reptiles. Here we describe Underwoodisaurus seorsus sp. nov., a new species similar to U. milii, but differing in its relatively plain dorsal and head patterns with only sparsely scattered pale tubercles, a much more gracile build, including longer snout, limbs and digits, smaller and more numerous fine scales on the dorsum, and the enlarged tubercles on the tail tending not to form transverse rows. The new species is known from few specimens and has only been encountered at mid elevations in the Hamersley Ranges, widely separated from the closest populations of U. milii in the northern Goldfields and Shark Bay in Western Australia. Given its rarity and small (potentially relictual) distribution this species may be of conservation concern.
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... This should mean that their publications may be treated more objectively by later herpetologists and their sensible and obvious taxonomic judgements widely used. Doughty and Oliver (2011) described as a new species, Underwoodisaurus seorsus as member of the U. milii complex from the Pilbara region of Western Australia, but significantly did not do a thorough review of the complex, as this would have necessitated the resurrection of the (until now) ignored Wells and Wellington taxon U. husbandi as a valid species-level taxon. Doughty and Oliver form part of a strongly anti-Wells and Wellington group known as the Wüster gang (se Hoser 2015a-f). ...
... The absence of East Australian U. milii from the phylogeny is conspicuous, but with the use of specimens from most other parts of the range, their phylogeny, provides strong molecular evidence for the recognition of the three species identified by Wells and Wellington (1985), these being U. asper, U. husbandi and U. milii and with these three being the minimum possible number of species in the species complex (and excluding the other later named taxon Underwoodisaurus seorsus) and that these are also the appropriate available names to be used. Of note is that if one factors in the East Coast U. milii as a separate taxon to the others, based solely on significant morphological divergence, the molecular results of Oliver and Bauer (2011) indicates that there are at least five separate lineages requiring taxonomic recognition, in addition to the species formally described by Doughty and Oliver (2011), giving a total count for the currently (treated by most as) monotypic genus Underwoodisaurus of 6 full species. Noting that the relevant populations can easily be distinguished on a morphological basis, I therefore name as new species within the U. milii complex, the currently unnamed divergent lineages from southern Australia (Victoria, southern inland NSW to the SA/WA border area) and that form from south-west Western Australia near Perth, while recognizing the other named forms (as listed by Wells and Wellington 1985 and that described by Doughty and Oliver 2011) as full species at the same taxonomic level. ...
... Of note is that if one factors in the East Coast U. milii as a separate taxon to the others, based solely on significant morphological divergence, the molecular results of Oliver and Bauer (2011) indicates that there are at least five separate lineages requiring taxonomic recognition, in addition to the species formally described by Doughty and Oliver (2011), giving a total count for the currently (treated by most as) monotypic genus Underwoodisaurus of 6 full species. Noting that the relevant populations can easily be distinguished on a morphological basis, I therefore name as new species within the U. milii complex, the currently unnamed divergent lineages from southern Australia (Victoria, southern inland NSW to the SA/WA border area) and that form from south-west Western Australia near Perth, while recognizing the other named forms (as listed by Wells and Wellington 1985 and that described by Doughty and Oliver 2011) as full species at the same taxonomic level. This overdue revision of the taxonomy of the U. milii group allows scientists to better quantify what taxa they may be studying by referring them to the appropriate species, especially noting that many potential differences between populations are not known at this stage. ...
Numerous studies and reclassifications of the Australian gecko family Carphodactylidae have been published in the previous three decades. These have resulted in the publication of a significant body of data, leading to the recognition of new genera and species. Molecular studies have indicated further unnamed groups at both generic and species levels. These taxa are all also readily identifiable on the basis of morphology.
The obvious unnamed taxa have therefore been formally described and named according to the rules of the
International Code of Zoological Nomenclature (Ride et al. 1999).
The genus Saltuarius Couper, Covacevich and Moritz, 1993 is divided along obvious phylogenetic and
morphological lines into two, the new genus being named Shireengecko gen. nov.. The two genera are both
split into two subgenera.
Phyllurus Schinz, 1822 is split four ways with the three new genera being Oxygecko gen. nov., Couperus gen.
nov. and Teesgecko gen. nov. respectively.
Two divergent species groups of Knob-tailed Gecko Nephrurus Günther, 1876 sensu lato are formally named
herein as subgenera, as Quazinephrurus subgen. nov. and Paranephrurus subgen. nov..
Nine new species are also formally named.
These are two within the genus Saltuarius sensu stricto as defined in this paper; one within Uvidicolus Oliver
and Bauer, 2011; one within Carphodactylus Günther, 1897; one within Nephrurus sensu stricto as defined in
this paper; two smooth knob tailed geckos (Quazinephrurus subgen. nov.) and two species of Thick-tailed
gecko Underwoodisaurus Wermuth, 1965. There are also four subspecies formally named.
The first ever key to the six species of Underwoodisaurus as defined herein is provided.
Carphodactylidae are also divided into four obvious tribes, with a further two subtribes identified and named.
Keywords: Taxonomy; lizards; gecko; nomenclature; Australia; Queensland; Cape York; Granite Belt; Wet
Tropics; New South Wales; South Australia; Victoria; Western Australia; Northern Territory; Carphodactylidae;
Carphodactylini; knob-tailed gecko; leaf-tailed gecko; thick-tailed gecko; Carphodactylus; Nephrurus; Orraya;
Phyllurus; Saltuarius; Uvidicolus; Underwoodisaurus; new genus; Shireengecko; Oxygecko; Couperus;
Teesgecko; new subgenus; Quazinephrurus; Paranephrurus; Quazisaltuarius; Quazishireengecko; new
species; hoserae; adelynae; jackyae; covacevichae; blacki; coreyrentoni; ianrentoni; mensforthi; perthensis;
new subspecies; martinekae; bulliardi; kimberleyae; saxacola; new tribe; Carphodactylini; Shireengeckiini;
Nephruriini; Orrayini; new subtribe; Uvidicolina; Oxygeckoina; Nephruriina; Shireengeckiina.
... The distribution of C. pilbarensis sp. nov. is centred on the Pilbara craton and its description brings the total number of gecko species occurring in the region to 39 (Wilson & Swan 2013;Cogger 2014;Oliver et al. 2014a;, including five endemic species and one subspecies, making it one of the most diverse regions for geckos in Australia (see also Powney et al. 2010;Doughty & Oliver 2011;Doughty et al. 2011b;Pepper et al. 2013b). Less well known, but emerging as an important centre is the North West Cape, which now has three described endemic taxa, all recognised in the last decade: C. tuberculatus sp. ...
We provide a taxonomic revision of the genus Crenadactylus, a group of very small clawless geckos from western and central Australia, with currently only one recognized species and four subspecies. Morphological comparisons were made on genotyped specimens from two recent genetic studies, then with an expanded sample to encompass all specimens to determine diagnostic characters in addition to morphological and geographic boundaries. Based on our findings, we elevate the subspecies Crenadactylus ocellatus ocellatus from south-Western Australia and C. o. horni from the Central Uplands to full species. Consultation of the types of Diplodactylus (= Crenadactylus) bilineatus indicate they are C. ocellatus based on a dorsal pattern with intermixing of dark and pale scales not shared with any other taxa; we therefore maintain synonymy of 'D.' bilineatus with 'D.' ocellatus. We describe three new Western Australian species formerly allocated to C. o. horni: C. occidentalis sp. nov. from the western coast, C. tuberculatus sp. nov. from the Cape Range and C. pilbarensis sp. nov. from the Pilbara region. To stabilize the Kimberley taxa, we also raise C. ocellatus rostralis and C. ocellatus naso, both monophyletic taxa from the monsoonal tropics, to full species, while acknowledging further work is required on the C. naso species complex. All new species treated here possess distinctive morphological characters to diagnose them, including enlarged dorsal tubercles in C. tuberculatus sp. nov., a single enlarged supranasal in C. horni and a single enlarged postmental in C. pilbarensis sp. nov. Pattern was relatively conserved among taxa, with highly contrasting dark and pale longitudinal lines, with the exception of the type species C. ocellatus that possesses intermixed dark and pale scales and ocelli. Crenadactylus species are separated by deep genetic divergences and are usually allopatrically distributed. This indicates that despite being Australia's smallest geckos, or possibly because of it, these diminutive lizards have a long history of localised persistence through major climatic changes over millions of years.
... The rocky Pilbara region in the western arid zone is a special case, with many new endemics described recently (e.g. Aplin et al. 2006;Doughty et al. 2010Doughty et al. , 2012Doughty & Oliver 2011; Maryan et al. ...
Lizards restricted to rocky habitats often comprise numerous deeply divergent lineages, reflecting the disjunct nature of their preferred habitat and the capacity of rocky habitats to function as evolutionary refugia. Here we review the systematics and diversity of the predominantly saxicoline Australian marbled velvet geckos (genus Oedura) in the Australian arid and semi-arid zones using newly-gathered morphological data and previously published genetic data. Earlier work showed that four largely allopatric and genetically divergent lineages are present: Western (Pilbara and Gascoyne regions), Gulf (west and south of the Gulf of Carpentaria), Central (central ranges) and Eastern (Cooper and Darling Basins). None of these four populations are conspecific with true O. marmorata, a seperate species complex that is restricted to the Top End region of the Northern Territory. Top End forms share a short, bulbous tail whereas the other four lineages treated here possess a long, tapering tail. Morphological differences among the arid and semi-arid lineages include smaller body size, tapering lamellae and a shorter tail for the Gulf population, and a partially divided rostral scale in the Western population compared to the Central and Eastern populations. Accordingly, we resurrect O. cincta de Vis from synonymy for the Central and Eastern lineages, and regard this species as being comprised of two evolutionary significant units. We also describe the Gulf and Western lineages as new species: Oedura bella sp. Nov. and O. fimbria sp. Nov., respectively. We note that a predominantly arboreal lineage (the Eastern lineage of O. cincta) is more widely distributed than the other lineages and is phylogenetically nested within a saxicoline clade, but tends to have a deeper head and shorter limbs, consistent with morphological variation observed in other lizard radiations including both saxicoline and arboreal taxa.
... The Pilbara bioregion of northwestern Australia is home to many endemic plants and animals, including Leeuwen's wattle (Acacia leeuweniana), the Pilbara barking gecko (Underwoodisaurus seorsus), and the Pilbara Ningaui (Ningaui timealeyi) (Maslin & van Leeuwen 2008;Gibson & McKenzie 2009;Doughty & Oliver 2011;Fig. 1). ...
Conservation decision tools based on cost-effectiveness analysis are used to assess threat management strategies for improving species persistence. These approaches rank alternative strategies by their benefit to cost ratio but may fail to identify the optimal sets of strategies to implement under limited budgets because they do not account for redundancies. We devised a multiobjective optimization approach in which the complementarity principle is applied to identify the sets of threat management strategies that protect the most species for any budget. We used our approach to prioritize threat management strategies for 53 species of conservation concern in the Pilbara, Australia. We followed a structured elicitation approach to collect information on the benefits and costs of implementing 17 different conservation strategies during a 3-day workshop with 49 stakeholders and experts in the biodiversity, conservation, and management of the Pilbara. We compared the performance of our complementarity priority threat management approach with a current cost-effectiveness ranking approach. A complementary set of 3 strategies: domestic herbivore management, fire management and research, and sanctuaries provided all species with >50% chance of persistence for $4.7 million/year over 20 years. Achieving the same result cost almost twice as much ($9.71 million/year) when strategies were selected by their cost-effectiveness ranks alone. Our results show that complementarity of management benefits has the potential to double the impact of priority threat management approaches.
... It is a relatively large gecko with a snout-vent length of about 80 mm [Cogger, 2000]. The second species of the genus, U. seorsus , was only recently described from the Pilbara region in Australia [Doughty and Oliver, 2011]. ...
Geckos (Gekkota) are a highly diversified group of lizards with an exceptional diversity in sex-determining systems. Despite this intriguing documented variability, data on sex determination in many lineages is still scarce. Here, we document the previously overlooked heteromorphic ZZ/ZW sex chromosomes in the thick-tailed gecko, Underwoodisaurus milii , a member of the ancient lineage of pygopodoid geckos. The finding of female heterogamety within pygopodoid geckos was unexpected, as until now only male heterogamety and environmental sex determination have been reported in this group, and female heterogamety was known only in distantly related gecko families separated from the pygopodoid geckos around 150 million years ago. The W chromosome in U. milii is highly heterochromatic and contains a large number of telomeric-like repeats comparable to around 50% of all telomeric-like sequences present in male genomes. The accumulation of these repeats might have been responsible for the considerable size expansion of theW chromosome in comparison to the Z chromosome. The heteromorphic ZZ/ZW sex chromosomes with accumulated telomeric-like repeats in the thick-tailed geckos further illustrate the exceptional diversity of sex-determining systems in geckos and add important information to our understanding of the evolution and phylogeny of sex-determining systems in reptiles.
... The Pilbara is known to harbour a suite of endemic vertebrates (How & Cooper, 2002; How & Dell, 2004; Gibson & McKenzie, 2009; Doughty et al., 2011a), with studies of widespread arid zone taxa typically revealing morphologically and genetically divergent Pilbara populations (e.g. Baverstock et al., 1983; Painter et al., 1995; Aplin & Donnellan, 1999; Blacket et al., 2000; Pepper et al., 2006; Ford & Johnson, 2007; Oliver et al., 2010; Doughty & Oliver, 2011; Melville et al., 2011). In addition, emerging results from fine-scale molecular studies of terrestrial vertebrates suggest substantial cryptic diversity and complex genetic patterns across the Pilbara landscape (Pepper et al., 2008Pepper et al., , 2011a Shoo et al., 2008; Doughty et al., 2010 Doughty et al., , 2011b Catullo et al., 2011). ...
We review the biogeography of the Pilbara, synthesize information on the geological and landscape history of this region and surrounds, and assess fine-scale genetic structure across multiple taxa to examine hypotheses concerning the distribution of genetic lineages. We use this to provide a baseline for future biological studies in an ancient area of endemism.
The Pilbara region, Western Australia.
Literature is summarized, including the history of Pilbara landscapes and climate, and previous biogeographical work. We used mitochondrial DNA phylogenetic datasets of seven co-distributed gecko (diplodactyline and gekkonine) lineages to assess the monophyly of Pilbara lineages, and concordance with geological and habitat divisions.
The Pilbara harbours taxa genetically distinct from their non-Pilbara relatives, despite close geographical proximity of populations. This is emphasized at the eastern and southern margins of the Pilbara, where habitat gradients are pronounced. In contrast, the northern margin, where sandy substrates of the Pilbara meet the dunes of the northern deserts, exhibits little genetic differentiation. Within the Pilbara, diversification patterns are idiosyncratic and may reflect species-specific ecological differences. However, a repeated north/south partitioning of genetic diversity is evident across taxa. An additional emerging pattern is an east/west genetic division in the northern Pilbara, which may relate to major drainage divides and geological discontinuities associated with east and west Pilbara terrains.
The Pilbara is an area of exceptionally high biotic diversity and endemism. The broader biogeographical patterns revealed in our molecular analyses are consistent with those recently identified using species richness patterns of invertebrates. Future studies of additional taxa using multiple molecular markers will provide the means to test and refine the biogeographical hypotheses presented here. Understanding the biogeography of the Pilbara and the partitioning of genetic diversity across the ancient and heterogeneous landscape is of paramount importance in the face of rapidly expanding economic and developmental pressures.
The gecko Diplodactylus savagei is restricted to the rocky Pilbara and Ashburton regions of Western Australia. Recent collections have enabled a reappraisal of morphological and genetic diversity within the taxon. Analysis of 1200 base pairs of the mtDNA gene ND2 and surrounding tRNA found strong support for three lineages within D. savagei: an eastern clade (which includes the type location of D. savagei from Marble Bar), a southern clade and a north-central clade. The eastern and southern clades did not differ in morphology or dorsal pattern. Although there are several subtle differences in morphological characters between the eastern and southern clades compared to the north-central form, there were clear differences in dorsal pattern with the north-central forms having finer, widely-scattered spots, a pale dorsal border to the loreal stripe and a gradual transition between the dorsal and ventral colouration. We describe the north-central form as a new species, D. galaxias sp. nov., based on the distinctiveness of its colour pattern, subtle morphological differences, mtDNA divergence and maintenance of these differences at the edge of the western Hamersley Range where the north-central and southern clades come into contact.
We describe a new species of Varanus similar to V. caudolineatus and V. gilleni but distinguishable from each of these taxa on genetic and morphological criteria. The three species are closely related and together constitute a species group within subgenus Odatria. The new species is restricted to the Pilbara region of Western Australia and appears to be sympatric with V. caudolineatus at several localities. It is more widely separated from known populations of V. gilleni. The new species is associated with mulga woodland and is at least partially arboreal, but little else is known of its ecology. Combined morphometric and meristic analyses indicate complex patterns of sexual dimorphism in all three species, including relative body elongation in females that is reflected in higher modal presacral vertebral counts in females than males of each species. Body elongation of females needs to be taken into account in future analyses of sexual dimorphism in varanid lizards.
The original illustration and descriptions of Phyllurus milii Bory de Saint Vincent, 1823 are most likely to be based on Nephrurus levis occidentalis Storr, 1963, not the species to which the name milii has been consistently applied for nearly 180 years. In the absence of types, a neotype (WAM R34085) is designated, representing the species to which the name milii has been applied by all subsequent authors, both for this species and Gekko dorrensis Peran, 1807. The senior name is identified as a nomen oblitum, and the junior name a nomen protectum, stabilising nomenclature of this species. The Thick-tailed or Barking Gecko, Nephurus milii (Bory de Saint-Vincent, 1823) is a widespread, large and readily identifiable species of southern Australia. Although the generic name has been subject to some debate, the species having been variously assigned to the genera or subgenera Phyllurus (e.g., Bory de Saint-Vincent, Kluge, 1991/ 1993), the species name has been consistently applied since 1823, although variously misspelt on occasion as nihi, miliusii, myliusii, milhus or milusii (see Bauer and Henle, 1994). Since 1934 (Loveridge, 1934), the specific epithet has mostly been correctly spelt. The species was named for Lieutenant-commander (later Baron) Pierre-Bemard Milius of Bordeaux, second-in-eommand of the corvette Naturaliste, one of two original ships of the French Baudin Expedition of 1800-1804 to Australian waters, with Bory de Saint-Vincent himself, also of Bordeaux, being one of two zoologists on the same ship (Comell, 1974). Bory de Saint-Vincent (1825, 1828) credited Milius with the discovery of the species, and for providing the colour plate drawn from life which accompanies the description. There is no evidence that Milius visited Australia, and more specifically the type locality (see below), other than with the Baudin Expedition. Bory de Saint-Vincent's description of this species was published in a popular encyclopedia of natural history several years after the premature death of Francois Peron (zoologist on the other ship of the Expedition, the Geographe) who had been preparing the zoological results of the Expedition for publication. The name appears three times in the encyclopedia, with a text account in volume 7 (Bory de Saint-Vincent, 1825: 183-184), another account in volume 13 (Bory de Saint-Vincent, 1828: 464-465), and a plate in the atlas, numbered the last volume in the series. Brygoo (1991) stated that the plate, although included in a volume with a title page date of 1831, was actually published as the first livraison of this volume in July 1823. Consequently, the plate validates the name, which must be regarded as published in 1823/ with the illustrated specimen the holotype. Two large gecko species of similar proportions co-exist in the Shark Bay region, including both Bemier and Dorre Islands: Nephrurus milii and Nephrurus levis occidentalis Storr, 1963 (Storr and Harold, 1978). As no type material can be identified (see below), the identity of the gecko named by Bory de Saint-Vincent must be based on his description and the accompanying plate, and primarily on the latter. Bory de Saint-Vincent (1825) stated: PHYLLURE DE MILIUS, Phyllurus Milii, N. (V. pI. de ce Dict.) Notre ancien et illustre ami le capitaine de vaisseau Milius, ci-devant gouvemeur de Mascareigne, maintenant charge du bonheur de la Guiane fran<;aise, a decouvert cette espece dans l'Australasie sur les rives de la baie des Chiens-Marins; nous lui en devons la figure et la description. Plus petite que la precedente, mais proportionellement plus haute sur jambes, sa h?te est obtuse, sa couleur d'un rouge de brique en dessus, qui ne permit que difficilement de la distinguer sur la terre rougeatre OU elle se tenait.
The concept of the skink lizard genus Eremiascincus Greer, 1979 is expanded to include eleven species [antoniorum, brongersmai, butlerorum, douglasi, emigrans, fasciolatus, isolepis, richardsonii, musivus sp. nov., pardalis, timorensis], eight of which [antoniorum, brongersmai, butlerorum, douglasi, emigrans, isolepis, pardalis, timorensis] (comb. nov.) previously belonged to Glaphyromorphus Wells & Wellington, 1983. This decision is based on the results of three recent studies, which indicated that ‘Glaphyromorphus’ was a polyphyletic assemblage representing a morphotype within Australian sphenomorphine skinks. In addition, we describe a new species of Eremiascincus based on morphological and molecular genetic evidence. The new species is distributed in coastal areas of the Pilbara region, Dampierland and the Great Sandy Desert in northwestern Western Australia. Eremiascincus musivus sp. nov. differs from regional congeners by possessing a characteristic dorsal pattern comprising numerous whitish and dark spots which align to form a diffuse reticulum, a pale vertebral stripe, more slender body and smaller body size, 52–62 paravertebral scales, scales along top of the fourth toe with oblique sutures on basal quarter to third of digit, subdigital lamellae of fourth toe undivided and feebly keeled and 10–15 plantar scales. The description of E. musivus sp. nov. brings the number of species of Australian Eremiascincus to seven.
We describe a new species of Varanus similar to V. caudolineatus and V. gilleni but distinguishable from each of these taxa on genetic and morphological criteria. The three species are closely related and together constitute a species group within subgenus Odatria. The new species is restricted to the Pilbara region of Western Australia and appears to be sympatric with V. caudolineatus at several localities. It is more widely separated from known populations of V. gilleni. The new species is associated with mulga woodland and is at least partially arboreal, but little else is known of its ecology. Combined morphometric and meristic analyses indicate complex patterns of sexual dimorphism in all three species, including relative body elongation in females that is reflected in higher modal presacral vertebral counts in females than males of each species. Body elongation of females needs to be taken into account in future analyses of sexual dimorphism in varanid lizards.
Native frog, mammal and reptile specimen data in the Western Australian Museum were examined from the western third of the Australian continent covering nearly 22 degrees of latitude and 16 degrees of longitude and encompassing tropical, desert and temperate regions. The timing of specimen data collection and collecting effort were evaluated and show that large areas of the State remain poorly sampled. The great majority of the collections have been made over the last 50 years and taxonomic status of many vertebrate species is still in review with several new species being described. Systematic surveys need to be undertaken to address the inadequacy of information on vertebrate fauna distributions over large tracts of the desert and pastoral areas of Western Australia. The distribution of taxa endemic to Western Australia, threatened and priority taxa as well as restricted-range endemic taxa were examined over equal areas based on the 1:250 000 map series that covers the western third of the Australian continent. Endemic taxa are focused in the south-west of the state and along the west coast, while restricted-range endemics are more frequently distributed along the west and northwestern coasts. Threatened and priority taxa show a similar pattern to that of endemic taxa. The similarity of areas across Western Australia, based on the composition of their vertebrate fauna, indicates that there are four broad regions corresponding to the tropical north, the mesic south-west, the semi-arid southwestern interior and the arid Pilbara and desert areas. Additionally, regional areas defined under the IBRA scheme were examined for the number of sampling locations, endemic taxa in the various faunal groups and the richness of taxa, recorded. The Pilbara bioregion, one of the best-sampled areas of the State, showed limited concordance between vertebrate taxa similarity in half-degree cells and subregionall boundaries and relatively high heterogeneity in vertebrate fauna distribution across the bioregion.
We generated a multi-locus phylogeny to test monophyly and distributional limits in Australian toadlets of the genus Up- eroleia from the western arid zone of Australia. The molecular data were used in combination with a detailed assessment of morphological variation and some data on call structure to complete a taxonomic revision of the species that occur in this region. Our work reveals the existence of not two but five species in the region. Uperoleia russelli is restricted to the Carnarvon and Gascoyne Regions south of the Pilbara. Uperoleia micromeles is distributed from the Tanami Desert through the Great Sandy Desert and along the northern edge of the Pilbara. Uperoleia talpa was previously believed to be a Fitzroyland region endemic but it is further distributed along Dampierland and into the Roebourne Plain. Uperoleia glan- dulosa is a larger species than previously described as well as a greater habitat generalist, inhabiting the rocky Pilbara region and the sandy region around Port Hedland. We also describe a new species, U. saxatilis sp. nov., endemic to the Pilbara craton.