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Systematics of small Gehyra (Squamata: Gekkonidae) of the southern Kimberley, Western Australia: Redescription of G. kimberleyi Börner & Schüttler, 1983 and description of a new restricted range species

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Ongoing fieldwork and molecular research continues to reveal that the monsoonal tropics of northern Australia contain more vertebrate species than currently recognised. Here we focus on two morphologically distinctive, yet unrecognised forms in the genus Gehyra from the southern Kimberley region and surrounding deserts. We base our descriptions on a combination of unpublished genetic data and a morphological examination of voucher specimens. We recognise and re-describe G. kimberleyi, a species with a broad distribution extending over most of the south-west Kimberley, across the Great Sandy Desert and into the far northern Pilbara. This species has been previously assigned to G. pilbara owing to its frequent occurrence on termite mounds and short snout, but can be distinguished from G. pilbara and other regionally sym-patric Gehyra by its moderate body size, moderate number of pre-cloacal pores in males (12-17) and aspects of dorsal colouration. We also describe G. girloorloo sp. nov., a small rock-dwelling species with a short snout, low number of pre-cloacal pores in males (8-11) and pinkish-grey dorsal colouration with alternating series of indistinct pale spots and irreg-ular transversely-aligned dark blotches. The new species appears to be restricted to a relatively small region of exposed limestone karst in the south-west Kimberley and is entirely circumscribed by morphologically similar congeners.
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Accepted by A. Bauer: 18 Mar. 2016; published: 29 Apr. 2016
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ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2016 Magnolia Press
Zootaxa 4107 (1): 049
064
http://www.mapress.com/j/zt/
Article
49
http://doi.org/10.11646/zootaxa.4107.1.2
http://zoobank.org/urn:lsid:zoobank.org:pub:487B9053-86A9-444C-8AC1-EB01F2F518C8
Systematics of small Gehyra (Squamata: Gekkonidae) of the southern Kimberley,
Western Australia: redescription of G. kimberleyi Börner & Schüttler, 1983 and
description of a new restricted range species
PAUL M. OLIVER
1,3
, GAYLEEN BOURKE
1
, RENAE C. PRATT
1
, PAUL DOUGHTY
2
& CRAIG MORITZ
1
1
Division of Evolution, Ecology and Genetics, Research School of Biology, and Centre for Biodiversity Analysis, The Australian
National University, Building 116, Daley Road, Acton, ACT, 2601, Australia
2
Department of Terrestrial Zoology, Western Australian Museum, 49 Kew Street, Welshpool, Western Australia, 6016, Australia
3
Corresponding author. E-mail: Paul.Oliver@anu.edu.au
Abstract
Ongoing fieldwork and molecular research continues to reveal that the monsoonal tropics of northern Australia contain
more vertebrate species than currently recognised. Here we focus on two morphologically distinctive, yet unrecognised
forms in the genus Gehyra from the southern Kimberley region and surrounding deserts. We base our descriptions on a
combination of unpublished genetic data and a morphological examination of voucher specimens. We recognise and re-
describe G. kimberleyi, a species with a broad distribution extending over most of the south-west Kimberley, across the
Great Sandy Desert and into the far northern Pilbara. This species has been previously assigned to G. pilbara owing to its
frequent occurrence on termite mounds and short snout, but can be distinguished from G. pilbara and other regionally sym-
patric Gehyra by its moderate body size, moderate number of pre-cloacal pores in males (12–17) and aspects of dorsal
colouration. We also describe G. girloorloo sp. nov., a small rock-dwelling species with a short snout, low number of pre-
cloacal pores in males (8–11) and pinkish-grey dorsal colouration with alternating series of indistinct pale spots and irreg-
ular transversely-aligned dark blotches. The new species appears to be restricted to a relatively small region of exposed
limestone karst in the south-west Kimberley and is entirely circumscribed by morphologically similar congeners.
Key words: Australian Monsoonal Tropics, biodiversity, endemism, gecko, limestone, lizard, short range endemic
Introduction
The Australian Monsoonal Tropics (AMT) has long been known within Australia for its high vertebrate diversity
(e.g. Cracraft 1991; Slatyer et al. 2007; Bowman et al. 2010; Powney et al. 2010). Nevertheless, ongoing surveys
and genetic analyses indicate that the AMT is home to many unrecognised vertebrate species (Potter et al. 2011;
Oliver et al. 2012, 2014a). For instance, at least eleven new or resurrected endemic vertebrate species have been
added to the faunal inventory of this region since the beginning of 2014: a frog (Catullo et al. 2014), seven lizards
(Oliver & Parkin 2014; Oliver et al. 2014a, b; Oliver et al. 2016), a snake (Maddock et al. 2015), a rock wallaby
(Potter et al. 2014), and a dasyurid marsupial (Aplin et al. 2015). Many other genetic lineages that may represent
further undescribed species have also been identified (e.g. Smith et al. 2011; Oliver et al. 2012, 2014c; Potter et al.
2012; Catullo & Keogh 2014). These new lineages include both widespread, apparently generalist, savannah taxa
(Catullo et al. 2014; Oliver et al. 2014a), and more restricted lineages, often associated with isolated and/or
geologically distinct rock formations (e.g. Hoskin et al. 2008; Doughty et al. 2009; Doughty 2011; Pepper et al.
2013; Oliver et al. 2014b, c).
Geckos from the genus Gehyra are among the most abundant lizards in the AMT, and are often seen on trees
and rocks when spotlighting at night. Here we focus on the taxonomic status of the smaller-bodied Gehyra from the
southern edge of the Kimberley region in north-west Australia (Fig. 1). This area lies at the border of the AMT and
the arid zone and coincides with the uplifted ranges of the King Leopold Range in the west, the Durack Range in
the east and other associated ranges such as the Oscar Range in the south-west (Tyler et al. 2012; Pepper & Keogh
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2014). The fauna reflects this overlap, with many widespread arid zone taxa occurring at the northern extremities
of their range, and other monsoonal taxa occurring at the southern edge of their distribution (e.g. for
mammals—McKenzie 1981; Start et al. 2012). A conspicuous geological feature of this area is a series of exposed
fossilised reefs (Tyler et al. 2012). These outcrops are not particularly extensive, and tend to be narrow (usually
less than a few kilometres wide) and low (maximum elevation usually less than 100 m above the surrounding
blacksoil plains; Goudie et al. 1990). These limestone karst outcrops have a complex and highly eroded
microtopography and provide an important local refuge from aridity and fire (Cameron 1992). Reflecting this they
have a high local endemism of snails (Cameron 1992), however, the first endemic vertebrate of the limestone
ranges was only recently described, a large-bodied Oedura gecko (Oliver et al. 2014b).
FIGURE 1. Distribution of Gehyra kimberleyi (circles) and G. girloorloo sp. nov. (diamonds) from the Kimberley and
surrounding regions of Western Australia; lighter regions are higher in elevation.
Only three species of small (snout-vent length [SVL] < 70 mm) Gehyra are currently recognised from the King
Leopold and nearby ranges in the south-west Kimberley. One form is part of a widespread species complex of
small saxicoline forms with well-defined dorsal spotting currently referred to as G. nana Storr, 1978. Unpublished
molecular data indicate G. nana sensu lato to be a complex of several morphologically similar forms (R. Pratt,
unpublished data). A recently-described small-bodied form—G. spheniscus Doughty, Palmer, Sistrom, Bauer &
Donnellan—occurs largely in the high rainfall region of the north-west Kimberley, but with outlying records in the
western King Leopold Range (Doughty et al. 2012). Lastly, existing collections and databases suggest the
occurrence of G. pilbara Mitchell, 1965 throughout this area (Storr et al. 1990; Wilson & Swan 2013; Atlas of
Living Australia 2015). However, morphological (Storr et al. 1990; pers. obs.) and molecular genetic data
(unpublished data) indicate that the northern records of G. ‘pilbara’ are a different taxon that can be distinguished
from most other small-bodied Gehyra in the southern Kimberley (including G. p i l ba r a senso stricto) by larger body
size, more subdigital lamellae, short and deep head and dorsal pattern of less defined pale and dark markings on a
pale to mid-tan background. This form is also somewhat of a habitat generalist—it usually occurs on trees and
termite mounds away from saxicoline habitats but is also occasionally collected from rocks. Based on its medium
size, colour pattern, distribution and often arboreal habits, we infer that this species is G. kimberleyi Börner &
Schüttler, 1983, described from a single subadult arboreal Gehyra from a locality near Derby (at the south-western
edge of the Kimberley). Some authorities have recognised the availability of this name in the past (e.g., Bauer
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SYSTEMATICS OF KIMBERLEY GEHYRA
1994; Kluge 2001; Uetz & Hošek 2015), yet no redescription or indication of its distribution based on examination
of specimens have been provided. As the taxon was named in a self-published journal the name has generally been
ignored or simply missed owing to its obscurity (e.g., their synonomy of G. purpurascens Storr to G. pilbara has
been universally ignored or rejected). The type material of this species is difficult to access—consultation with
Australian state authorities indicate no collection or export permits to Germany from Australia were issued, and
they are currently housed in a private residence in Germany (Uetz & Hošek 2015). However, although the short
original description and accompanying photographs indicate the holotype is a small sub-adult, it provides sufficient
images and diagnostic characters to assign the northern G. pilbara s.l. form to this name. Here we present a
redescription of this species based on much more extensive material in Australian museums.
Recent surveys have also revealed another distinctive form of Gehyra in the far southern Kimberley that
appears to be restricted to a small area of limestone karst just south-east of Fitzroy Crossing (Fig. 1). This small
(SVL < 50 mm) Gehyra is morphologically similar to the widespread complex of geckos currently referred to as G.
nana (small size, single egg, pattern of light and dark spots and blotches). The limestone form differs from the
other four species from this region in having small body size, no wedge of granules at the base of the digits
(diagnostic of G. spheniscus), a lower number of pre-cloacal pores in males and a pale pinkish dorsal colouration
with a pattern of alternating dark smudges and pale bars. Unpublished genetic data using mtDNA and nDNA
indicate the limestone form is highly divergent from G. kimberleyi and all G. nana complex forms. This genetic
data will be presented elsewhere, but as the limestone form is morphologically diagnosable, we describe it as a new
species below.
Material and methods
We examined recently-collected specimens held at the Australian National University (ANU, Canberra), and
specimens housed at the Western Australian Museum (WAM) or Museum Victoria (NMV), where all type material
is deposited (see type lists and Appendix). For detailed morphological comparisons, we chose specimens for which
matching molecular data was available (unpublished data), with the exclusion of immature or poorly-preserved
individuals. Additional comparative material of G. nana and G. pilbara were also examined (Appendix).
We examined 15 morphometric and 6 meristic characters (Table 1), comprising body, limb and head
proportions, head scalation, pre-cloacal pore counts and counts of divided lamellae beneath the fourth finger and
toe. Character descriptions generally follow those of Doughty et al. (2012). Bilateral measures were taken
unilaterally from the right side of the body, unless prevented by damage or poor preservation. Measurements were
taken to the nearest 0.1 mm using Mitutoyo electronic callipers and scale and pore counts were made under a
dissecting microscope (Leica MZ8). Descriptions of colouration in life were made based on photographs of a
subset of specimens taken either in life or just prior to preservation, in addition to notes taken in the field.
Taxonomy
Gehyra Gray, 1834
Type species. Gehyra pacifica Gray, 1834 (= Gecko oceanica Lesson, 1830 synonymy fide McCann 1955), by monotypy.
Gehyra kimberleyi Börner & Schüttler, 1983
Robust termitaria gecko
Figs. 2–4
Holotype. BSRC Geck 65 SC (private collection of Achim-Rüdiger Börner, Germany), ‘5 km south of Derby,
W.A.’
Diagnosis. Digits broadly expanded basally and subdigital scansors present on all digits of manus and pes.
Digit I of manus and pes clawless, penultimate phalynx of digits II–V free from scansorial pad. Differs from non-
Australian Gehyra by the combination of: absence of webbing between third and fourth toes, absence of a skin fold
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along the posterior hindlimb and medium adult size (SVL < 62 mm). Differs from Australian Gehyra by the
combination of moderate size (SVL < 62 mm), short snout (SnEye/HL mean 0.42) and deep head (HD/HL mean
0.48), subdigital lamellae divided without basal wedge of granules, moderate number of pre-cloacal pores (12–17),
postmentals not contacting the second infralabial, one pair of chin shields and dorsal background colour light
reddish-brown to pale tan or greyish-brown with diffuse, widely-spaced pale spots and small dark-brown blotches.
Details of holotype. From original description (in mm). SVL 36; TailL 39; HeadL (tip of rostral to anterior
corner of ear) 9; HW 7.3; HD 4.3; HD/SVL 0.119; SupLab* 6/7; InfLab* 6/7; FingSubDL 6; ToeSubDL 9; rostral
with cleft, undivided; first SupLab higher than second (*not defined, possibly measured to centre of eye).
Description of species. Range of variation for measurements and scale counts are presented in Table 1. A
medium-sized (adult SVL mean 50.4 mm, range 42.0–61.3 mm), moderately robust gecko. Head deep (HD/HL
0.48, 0.40–0.58), moderately wide (HW/HL 0.86, 0.76–0.97), widest posterior to eye, tapering to a rounded snout,
narrowing posteriorly to a slightly constricted neck. Snout short (SnEye/HL 0.42, 0.39–0.46), generally shorter
than twice eye length (OrbL/SnEye 0.58, 0.54–0.63), occasionally slightly concave on dorsal surface, canthus
rounded; in profile view, snout convex with blunt tip. Body moderately long (TrunkL/SVL 0.42, 0.37–0.47) and
robust, slightly depressed.
TABLE 1 . Measurements (mean [range], in mm) and meristic data (mode [range]) for the four small-bodied Gehyra
species treated here.
On head, dorsal scales largest on snout, ~2.5 times larger than scales on crown of head and body and slightly
projecting and imbricate. Nostrils large and circular, bordered by rostral, supranasal, two postnasals of similar size
and first supralabial; supranasals round to triangular in shape, usually separated by a single large internarial scale.
Rostral scale wider than high (RosH/RosW 0.64, 0.56–0.74), slightly gabled, usually with a deep medial cleft
~60% of rostral height. Supralabials 7–9 (mode 8), infralabials 7–8 (mode 7). First supralabial higher than
subsequent supralabials. Supralabials bordered dorsally by enlarged row of scales posteriorly that decrease in size
G. kimberleyi
N = 19
(M = 11/F = 8)
G. girloorloo sp. nov.
N = 17
(M = 10/F = 7)
G. nana (south Kimberley)
N = 40
(M = 20/F = 20)
G. pilbara
N = 14
(M = 4/F = 10)
SVL 50.4 (42.0–61.3) 42.9 (39.8–47.4) 40.0 (35.0–44.3) 41.4 (33.4–45.1)
HL 12.0 (9.9–14.1) 10.4 (9.7–11.5) 10.3 (8.9–11.9) 9.4 (7.9–10.6)
HW 10.4 (8.5–12.6) 8.6 (7.6–9.1) 8.4 (7.6–9.4) 8.0 (6.9–9.1)
HD 5.8 (4.5–7.9) 4.7 (3.8–5.6) 4.3 (3.6–5.1) 5.2 (4.5–6.4)
ArmL 5.5 (4.1–7.7) 4.9 (4.6–5.6) 4.4 (3.6–5.8) 4.4 (3.0–5.3)
HindL 6.0 (4.6–8.7) 4.9 (4.4–5.5) 4.8 (3.6–6.7) 4.5 (3.1–5.6)
TrunkL 21.1 (16.2–27.9) 17.8 (15.1–20.7) 15.8 (12.0–19.5) 17.3 (15.1–19.6)
NarEye 4.0 (3.2–5.2) 3.4 (3.0–3.8) 3.4 (2.9–4.0) 2.9 (2.4–3.4)
SnEye 5.0 (4.1–5.9 ) 4.4 (3.9–4.9) 4.4 (3.7–5.0) 3.9 (3.6–4.3)
EyeEar 3.5 (2.8–4.2) 2.8 (2.3–3.2) 2.9 (2.4–3.9) 2.5 (1.9–3.2)
InterN 1.5 (1.1–2.1) 1.3 (1.2–1.5) 1.3 (0.9–1.6) 1.1 (0.9–1.3)
InterOrb 3.6 (3.0–4.4) 2.9 (2.5–3.3) 3.0 (2.5–3.7) 2.8 (2.3–3.1)
OrbL 2.9 (2.4–3.4) 2.6 (2.3–2.9) 2.3 (1.6–2.8) 2.3 (2.0–2.6)
RosH 1.1 (0.9–1.4) 0.9 (0.7–1.1) 1.0 (0.6–1.3) 1.0 (0.8–1.3)
RosW 1.8 (1.4–2.1) 1.5 (1.4–1.8) 1.6 (1.3–2.0) 1.5 (1.2–1.9)
SupLab 8 (7–9) 8 (7–9) 8 (7–10) 7 (7–8)
InfLab 7 (7–8) 7 (7–9) 8 (6–9) 7 (7–8)
InterNsc 1 (0–2) 0 (0–2) 1 (0–1) 0 (0–1)
PCP 16 (12–17) 9 (8–11) 14 (11–17) 13 (13–14)
FingSubDL 6 (5–7) 6 (5–6) 5 (5–6) 5 (5–6)
ToeSubDL 7 (6–9) 6 (6–7) 6 (5–7) 6 (5–7)
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as for supralabials. Mental scale roughly triangular, divides postmentals 20–40%. Postmentals in contact with first
infralabial only; single pair of outer chin shields smaller with rounded outer edge, ~two-thirds the height of
postmentals, in contact with first and second infralabials, thereby excluding postmental from contact with second
infralabial. Second or third infralabial notched and in contact with first parinfralabial. Eye small (OrbL/HL 0.24,
0.21–0.27), circular, small fringe of mostly rounded scales extends from the postero-dorsal to the anterio-ventral
edge. Ear openings moderately large and circular.
FIGURE 2. Preserved specimens of Gehyra kimberleyi, showing variation in dorsal patterns. Scale bar = 1 cm.
Dorsal scales homogeneous, small, rounded and juxtaposed; gular scales granular; ventral midbody scales 2–3
as large as dorsal scales, slightly imbricate and tending to be diamond-shaped; limbs covered in granular scales,
scales on anterior surfaces enlarged, especially on legs. Pre-cloacal pores 12–17, mode 16, perpendicular to body
distally and medial curving forwards to form a sharply pointed ‘v’ with anteriorly-directed apex. Usually a single
small post-cloacal spur present on each side in males. Females lack pre-cloacal pores and prominent post-cloacal
spurs.
Limbs very short (ArmL/SVL 0.11, 0.09–0.13; HindL/SVL 0.12, 0.10–0.17); five digits on each limb, claws
present on digits 2–5, claws long and free, arising from dorsal surface of expanded toe pads and extending above
and well beyond toe pad, row of elongate and sharply pointed distal scales on periphery of dorsal surface of toe; toe
pads ovoid; subdigital lamellae divided, 5–7 (mode 6) pairs on fourth finger, and 6–9 (mode 7) on fourth toe, apical
lamellae wedge-shaped.
Tail moderately long and moderately thin to slightly swollen, slightly dorso-ventrally flattened in cross-
section, tapering to fine tip. Dorsal scales of tail flattened with rounded posterior edge, tending to be arranged in
rows. Enlarged, wide series of scales beneath tail, bordered peripherally by 2–4 rows of enlarged scales, decreasing
in size on lateral surfaces.
Colouration in preservative. Background dorsal colour varies from dark reddish- or greyish-brown with fine
dark stippling to light salmon-pink, pale and patternless in larger individuals (Fig. 2). Dorsal surfaces of head, torso
and tail usually with scattered poorly-defined small, dark greyish-brown spots, blotches, flecks or short bars; pale
grey spots interspersed among the darker markings, usually forming rows of 4–6 spots; on the neck and crown,
blotches are usually separated, on the torso often transversely aligned to form loosely-defined bands on the body
and on the tail often forming concentric bands. Dark longitudinal streaks on lateral surface of neck posterior to eye
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or a series of small dark blotches. Limbs with similar pattern to dorsum, but generally less well defined. Ventral
colour pale, lightly stippled but more darkly pigmented anteriorly, especially in gular region. Iris pale silvery blue,
pupil vertical and pale with crenelated edge. Original tails with alternating dark and pale concentric rings or series
of blotches. Regenerated tails with no clear pattern, although indistinct longitudinal brown streaks are sometimes
apparent.
Colouration in life. Based on photographs of specimens captured in the south-western Kimberley. The dorsal
background colour is pale or golden tan to medium dark brown, and the dorsal pattern consists of scattered dark
brown blotches and smaller pale spots that occur in alternating rows, especially the vertebral zone, with smaller spots
occurring on the head, flanks and limbs. The dorsal pattern tends to be less distinct in larger specimens (Fig. 3).
FIGURE 3. Gehyra kimberleyi in life. Top: weakly marked adult specimen from Roebuck Plains Homestead; bottom: strongly
marked juvenile specimen from Broome (photographs—B. Maryan).
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Habitat and reproduction. Habitat notes from collectors of specimens held at the WAM indicate most were
collected from termite mounds (128 of 227 such records). Other collection locations indicate generalist habits, with
specimens collected from trees, under logs and other ground cover and including vertical rock faces. Like other
species in the Gehyra variegata-punctata species-group (Doughty 1996), G. kimberleyi lays a single egg.
Distribution. Occurs throughout the southern Kimberley region, from Cape Leveque at the northern edge of
the Dampier Peninsula in the west, as far north as Mt Elizabeth in the central Kimberley, and north-east to Lake
Argyle near Kununurra (Fig. 1). Most records are from south of the King Leopold and Durack barrier ranges, with
several records south of the Fitzroy Crossing area. There are no records north of the Durack Range in the eastern
Kimberley, although this region is the most poorly surveyed of the Kimberley. In the Northern Territory, the species
is recorded from Buchanan Station. There are records of genotyped specimens as far south as the northern Pilbara
(15 km north-east of Nifty Mine and Meentheena Nature Reserve), with several specimens from along the 80 mile
beach extension of Dampierland between the west Kimberley and Pilbara regions. In the Great Sandy Desert, there
are records from Dragon Tree Soak Nature Reserve in the south to records further north. The Great Sandy Desert
remains poorly surveyed and this species may be more widely distributed in this region.
Etymology. Although the original description mentions the ‘Kimberleys’ (sic) (Börner & Schüttler 1983, p. 1),
no explicit etymology was provided. The presumed correct formation of the specific name is ‘kimberleyensis’,
which means the taxon is from the Kimberley region. The proposed name ‘kimberleyi’ would refer to a male
named Kimberley. The error, however, cannot be emended under current ICZN rules of nomenclature. The
suggested common name refers to species’ body habitus, and ‘termitaria’ indicates this species’ habitat preference
(although it is not exclusive to termitaria).
Comparison with other species. Gehyra kimberleyi can be distinguished from all non-Australian Gehyra by
the combination of absence of webbing between the third and fourth toes (versus present), the absence of a skin
fold along the posterior hindlimb (versus present) and its smaller size (max SVL < 65 mm versus > 65 mm).
Within Australia, G. kimberleyi can be distinguished from all members of the G. australis species-group (which
are largely restricted to the AMT) by its smaller size (max SVL < 65 mm versus max SVL > 65 mm), divided
subdigital lamellae (versus at least some undivided) and single egg per clutch (versus two).
Gehyra kimberleyi can be distinguished from most other members of the G. variegata-punctata group that also
occur in the AMT as follows; from G. xenopus Storr and G. spheniscus by the absence of a wedge of granules
between proximal lamellae (versus present), and in the case of the former species, smaller size (max SVL 62 versus
79 mm); from G. occidentalis Storr by its smaller body size (max SVL 62 versus 76 mm), deeper head, postmentals
3 x longer than wide (versus 4 x), lower number of subdigital lamellae (6–8 versus 7–10), fewer pores in males
(12–17 versus 18–30) and more clearly-defined spots on the dorsum; and from G. multiporosa Doughty, Palmer,
Sistrom, Bauer & Donnellan, 2012 by fewer pores in adult males (12–17 versus 20–49), dark markings on dorsum
not tending to form transverse bars and the absence of dark lateral head streaks.
Gehyra kimberleyi can be distinguished from both geographically proximate and type G. nana by short snout
and deep head, tan or greyish brown dorsal colouration (versus reddish), generally larger body size (SVL 42.0–61.3
versus 35.0–44.3), more larger, irregular and pale blotches (versus small and clearly defined), diffuse and
transversely oriented dark brown blotches on dorsum (versus clearly defined brown blotches or spots).
Gehyra kimberleyi overlaps with a number of other members of the G. variegata-punctata species-group from
the Australian arid zone, especially in the northern Pilbara. It differs from species in this group in having dorsal
markings of dark blotches that tend to coalesce to form transverse bars (versus continuous longitudinal lines or
network in G. variegata and G. montium—Hutchinson et al. 2014), and by having 7–9 supralabial scales (versus
8–10 in G. variegata and 8–11 in G. montium—Hutchinson et al. 2014). It is distinguished from members of the
Gehyra punctata species complex by its diffuse dorsal colouration (versus well defined dark and light spots or
blotches) and postmentals that do not contact the second supralabial.
From Gehyra pilbara (with which it has frequently been confused) it can be distinguished by its larger size
(max SVL 62.3 versus 45.1 mm), tall, narrow postmentals (versus extremely short and wide; cf. Fig. 4), and tan or
greyish-brown dorsal colouration (vs. reddish-brown).
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Gehyra girloorloo sp. nov.
Kimberley Karst gecko
Figs. 4–7
Holotype. WAM R175045 (field number CCM3257) (male), Gogo Station, Pillara Range, south entrance of
Menyous Gap (18.40439°S; 125.83698°E), collected on 4 November 2014 by P.M. Oliver, G. Armstrong and P.
Skipwith.
Paratypes. WAM R113727 (female), Cadjbut Mine 90 km south-east Fitzroy Crossing (18.7500°S;
126.1500°E); NMV D77029 and NMV D77030 (females), and NMV D77031 and NMV D77032 (males),
Ngumpan Cliff area (18.75625°S; 126.06474°E); WAM R175037 (PMO147) and WAM R175038 (PMO149)
(males), < 1 km south of Galeru Gorge (18.61407°S; 126.08386°E); WAM R175039 (CCM3242) and WAM
R175040 (CCM3243) (males), Gogo Station, Limestone Billy Hills (18.32724°S; 125.76498°E); WAM R175043
(CCM3246) (female), Gogo Station, Limestone Billy Hills (18.33516°S; 125.75244°E).
Diagnosis. Digits broadly expanded basally and subdigital scansors present on all digits of manus and pes.
Digit I of manus and pes clawless, penultimate phalanx of digits II–V free from scansorial pad. Differs from non-
Australian Gehyra by the combination of: absence of webbing between third and fourth toes, absence of a skin fold
along the posterior hindlimb and small adult size (SVL < 48 mm). Differs from all other Australian Gehyra by the
combination of small body size (SVL < 48 mm), divided subdigital lamellae without basal wedge of granules, short
snout and large eyes (OrbL/SnEye 0.58, 0.51–0.72), low number of pre-cloacal pores in males (8–11), postmentals
not in contact (at most in point contact) with the second supralabial, one pair of outer chin shields and pinkish-
brown dorsal background colour with pattern consisting of alternating bands of indistinct off-white spots and dark-
brown transverse blotches on heavily stippled background.
Description of holotype. Adult male with following dimensions (in mm): SVL 41.3, HD 4.7, HL 10.2, HW
8.2, HindL 5.4, ArmL 5.0, TrunkL 18.4, NarEye 3.2, SnEye 4.5, InterN 1.5, InterO 3.2, OrbL 2.4, RosW 1.5, RosH
0.9. Summary meristic data are as follows: SupLab 8; InfLab 7; InterNSc 0; PreCloacal 9; FingSubDL 6;
ToeSubDL 7.
FIGURE 4. Chin shield configuration in Gehyra treated here; A) G. kimberleyi WAMR175035, B) G. pilbara WAMR108632,
C) G. girlooloo sp. nov. WAMR175045 and D) G. nana WAMR175063. Scale bar = 1 mm.
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FIGURE 5. Dorsal and ventral views of the Gehyra girloorloo sp. nov. holotype (WAM R175045, CCM3257). Scale bar = 1
cm.
A small (41.3 mm SVL), slender gecko (Fig. 5). Head slightly depressed (HD/HL 0.46), moderately wide
(HW/HL 0.80), of similar width to midbody, widest at posterior edge of jaw, tapering to snout, narrows posteriorly
to a moderately constricted neck. Snout short (SnEye/HL 0.44), less than twice eye length (OrbL/SnEye 0.54),
rounded in dorsal profile, slightly convex from mid-eye to naris when viewed laterally in profile. Eyes moderately
sized (OrbL/HL 0.24), pupil vertical. Ear openings small and circular. Nostrils rounded, contacted by rostral,
supranasal, two postnasals and first supralabial; second (ventral) postnasal ~2 times larger than first (dorsal),
supranasals in contact, no internarial scale. Rostral scale rectangular with nearly flat dorsal edge, wider than high,
1.5 mm wide, 0.9 mm high (RosH/RosW 0.58), deeply furrowed, rostral crease extending from dorsal edge ~40%
of rostral height. Supralabials 8, infralabials 7. Scales on snout large, up to 3 times larger than scales on crown of
head and body, imbricate and slightly projecting along posterior edge; fold of skin across snout from eye to eye
(artefact of preservation). A row of slightly enlarged scales above supralabials gradually decrease in size
posteriorly. Mental scale triangular, divides postmentals ~35%; postmentals narrowly separated from second
infralabial on right side, in point contact on left side; single pair of outer chin shields with curved dorso-postero
edge, ~two-thirds the height of postmentals and ~30–40% smaller in area; outer chin shield in point contact with
first infralabial on right side, narrowly in contact on left.
Body long (TrunkL/SVL 0.46) and slender, slightly depressed and with a weak ventro-lateral fold posterior to
axilla. Dorsal midbody scales small and granular, relatively homogenous, conical, juxtaposed; lateral scales
slightly larger, more projecting and heterogeneous than those on dorsal surface, slightly imbricate; ventral scales
~3 times larger than dorsal scales, flat, diamond-shaped with rounded edges, juxtaposed. Pre-cloacal pores in a
series of 9, forming a short continuous chevron with apex orientated anteriorly; pores centred in scales, innermost
scales the largest. Two enlarged, rounded and protruding cloacal spurs present on either side of cloaca.
Limbs short (ArmL/SVL 0.12; HindL/SVL 0.13); dorsal and ventral scales generally small, rounded and
juxtaposed; scales on anterior edge of limbs slightly larger and imbricate. Fingers and toes 5; expanded toe pads
oval; distal row of dorsal scales on toe pads elongate and sharply pointed, forming a serrated ‘fringe’. Claws long
and free, present on digits 2–5, arising from dorsal surface of expanded toe pads and extending above and beyond
toe pad, subdigital lamellae divided, 6 pairs on fourth finger, 7 pairs on fourth toe, all pairs of lamellae in contact,
apical lamellae undivided and wedge-shaped.
Most of tail is original (52.0 mm, TailL/SVL 1.3), very thin, tapering gently to fine tip, slightly dorso-ventrally
flattened at base, proximal 31 mm section original, distal 21 mm section regrown. At widest part near base—4.3
mm wide, 3.3 mm deep. Dorsal caudal scales small, granular and relatively homogeneous, subcaudal scales in a
single large transversely widened series.
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Colouration of holotype (in preservative). Background colour pale grey, lightly stippled with darker gray, a
series of pale circular spots tending to form oblique transverse rows, enclosed by more extensive transversely-
oriented darker grey markings or larger variegations; crown of head with small pale spots above tympanum region,
snout and sides of head dark grey, no streaks emanating posteriorly from eye, labials heavily stippled; limbs as for
dorsum with smaller pale spots; undersurfaces pale off-white; dorsal surface of tail encircled by pale (wider) and
dark brown (thinner) bands.
Vari a t i o n . Table 1 presents ranges of variation for the characters measured. In a small number of specimens,
the postmental and second infralabial are in point contact, otherwise they are separated. The same pattern holds for
internasals—in most specimens the supranasals are in broad contact above the rostral, in a small number a tiny
internasal scale is present.
Colouration in preservative is as follows: background dorsal colour pale tan or grey (Fig. 6), densely stippled
with fine blackish-brown variegations, moderate sized (0.51.5 mm wide) poorly defined pale circular to oval spots
scattered on the dorsum, separated by concentrated patches of dark-brown markings that coalesce to form weakly
defined transverse bands; density and distribution of dorsal pattern elements varies along body: usually little or no
clear pattern on anterior portion of head, more clearly defined on nape with scattered small light spots and
indistinct dark blotches, and consisting of alternating series of comparatively large and more well defined pale
spots and indistinctly defined transverse brown bands on the torso and original tail. Limbs as for torso, with
scattered distinct to indistinct small pale spots; undersurfaces largely unpigmented, but with dense fine dark brown
maculations around the ventrolateral regions of head, limbs and ventral tail scales. Regrown tail light greyish-
brown with diffuse longitudinally aligned narrow dark streaks.
Colouration in life. The following notes are based on photographs of specimens NMV D77030 and WAM
R175037 shown in Fig. 7 and WAM R175042–3 (not shown). Dorsal surfaces pale purplish buff very finely
stippled with medium brown, overlain on dorsum and tail with alternating transverse series of pale spots lacking
borders and irregular dark-brown blotches that tend to be transversely aligned and intermittently bordered with
yellow-orange. Head with smaller pale and dark spots with scattered yellow-orange flecks or fine spots.
Regenerated tails often have a distinctive yellowish wash and very fine brown longitudinal markings. Iris dark
golden brown with fine black reticulations.
Habitat and reproduction. This species is closely associated with dissected limestone outcrops, and also
occurs on small trees and shrubs (Fig. 8). It tends to be less common on exposed vertical rock faces than its much
larger sympatric congener G. k o i r a Horner. The locally occuring form of Gehyra nana tends to utilise different
habitats—it is rarely found on limestone and generally on smaller boulders and rocky debris in surrounding areas.
Gehyra girloorloo sp. nov. can be abundant in its preferred habitat, for example, most large shrubs along an open
rock face in Menyous Gap had at least one specimen, often positioned head down close to the ground in what
appeared to be a hunting posture. In contrast, we found this species to be scarce in early July 2014 when
temperatures were low (< 15°C), with only two specimens observed over two nights of searching around Mt Piere
Station.
Like all Gehyra in the variegata-punctata species-group, this species lays a single egg. Some females were
gravid in November in spring at the end of the dry season.
Distribution. Gehyra girloorloo sp. nov. is only known from a small area of limestone outcropping in the
King Leopold Range of the south-west Kimberley on Gogo and Mt Piere Stations (Fig. 1). Known localities extend
in a narrow band from the limestone Billy Hills in the north-west, to just north-west of Ngumpan Cliff in the south-
east, a straight line distance of approximately 55 km.
Etymology. Girloorloo, pronounced “gir-loor-loo”, is a word the local Gooniyandi mob use for the limestone
this species appears to be restricted to. This species name was suggested by elders from the Gooniyandi mob who
speak for country on which this species occurs.
Comparisons. Gehyra girloorloo sp. nov. can be distinguished from all non-Australian Gehyra by the absence
of webbing between third and fourth toes (versus present), the absence of a skin fold along the posterior hindlimb
(versus present) and its generally smaller size (max SVL < 50 mm versus > 50 mm).
Within Australia, G. girloorloo sp. nov. can be distinguished from all members of the G. australis species-
group (which are largely restricted to northern Australia) by its smaller size (max SVL < 50 mm versus > 50 mm),
divided subdigital lamellae (versus at least some undivided), lower number of pores in the males ( < 12 versus >
12) and females laying just one egg per clutch.
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FIGURE 6. Small saxicoline Gehyra of the southern Kimberley, Western Australia. Top: G. girloorloo sp. nov. NMV D77030
near Ngumpan Cliff; middle: G. girloorloo sp. nov. WAM R175037 (PMO147), Galeru Gorge; bottom: G. nana NMV D77035
near Ngumpan Cliff (photographs—P. Horner and T. Parkin).
Gehyra girloorloo sp. nov. can be distinguished from members of the G. variegata group from the Australian
arid zone by its lower number of pre-cloacal pores in males (< 12 [mode 9] versus usually more than 10
[Hutchinson et al. 2014]), no conspicuous dark streaks emanating from behind the eyes, at most only light red hues
(versus rich reddish-brown) and less contrasting dorsal pattern combining diffuse dark markings (versus extensive
reticulations) and diffuse light spots (versus tending towards smaller and well defined). A further species in this
group, G. pilbara, is similarly small, but differs in dorsal colouration (reddish-brown vs. pinkish-grey), the presence
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of obviously enlarged loreal scales above the infralabials (versus absent) and in having an extremely short snout
resulting in short, wide postmentals (versus tall and thin in G. girloorloo sp. nov.) (Fig 4.).
FIGURE 7. Gehyra girloorloo sp. nov. paratypes showing variation in colour and pattern (in preservative). Scale bar = 1 cm.
FIGURE 8. Habitat of Gehyra kimberleyi (small trees in foreground) and G. girloorloo sp. nov. (karstic limestone in
background) at Gogo Station, Western Australia (photograph—P.M. Oliver).
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Gehyra girloorloo sp. nov. can be distinguished from most other members of the G. variegata group that also
occur in the AMT as follows: from G. xenopus and G. spheniscus by the absence of a wedge of granules between
proximal lamellae (versus present), and in the case of the former species, also much smaller size (max SVL 48
versus 79 mm); from G. occidentalis by its lower number of subdigital lamellae (5–7 versus 7–10) and small body
size (max SVL 48 mm versus 76 mm); and from G. multiporosa by the absence of dark lateral head streaks, and
fewer pores in males (8–11 versus 20–49).
A final taxon from this region, G. nana, is a complex of species (unpublished data). However, G. girloorloo sp.
nov. can be distinguished from both geographically proximate and type G. nana by its pinkish-grey dorsal
colouration (versus reddish), background stippled (versus plain), larger and more diffuse pale spots (versus small
and clearly defined), diffuse and transversely-oriented dark brown blotches on dorsum (versus clearly defined
brown blotches or [usually] spots), and low number of pre-cloacal pores in males (8–11 versus 11–17). Gehyra
nana from around localities from where G. girloorloo sp. nov. has been recorded are also particularly small (SVL
39.0, 35.0–41.9 mm) and strongly spotted (see Fig. 5).
Discussion
The mesic and topographically complex ranges and islands of the northern Kimberley are widely recognised for
their high biodiversity and endemism, and attract considerable scientific interest (Burbidge and McKenzie 1978;
Palmer et al. 2013). In contrast, the low ranges fringing the southern Kimberley are not currently recognised as an
endemism hotspot, with the first endemic vertebrate species only recently described (Oliver et al. 2014b). The
distribution of G. girloorloo sp. nov. suggests that it may also be restricted to the limestone outcrops of the southern
Kimberley. Ongoing work in other gecko lineages indicates that additional taxa from these limestone ranges remain
unrecognised (R. Pratt, R. Laver, P.M. Oliver submitted).
Gehyra girloorloo sp. nov. is sympatric with the much larger saxicoline congener G. k oi r a . It is also regionally
sympatric with G. nana (predominantly saxicoline), G. kimberleyi (termite mounds, otherwise generalist) and G.
australis (arboreal). However, although the range of the latter three species seems to largely circumscribe that of G.
girloorloo sp. nov., and all have been recorded within less than a kilometre of the new species, they have not yet
been observed using the same limestone outcrops. Of particular note is the absence of the small and widespread
saxicoline generalist G. nana from limestone inhabited by G. girloorloo sp. nov.—even though this species often
utilises narrow gaps on and around rock boulders, and has been found on laterites just metres from limestone
outcrops inhabited by G. girloorloo sp. nov. This partitioning of saxicoline environments suggests that variation in
microhabitats in outcrops of differing geology is mediating co-existence in regional sympatry, presumably on the
basis of varying physiological tolerances and/or competitive ability.
Acknowledgements
We thank the elders from the Gooniyandi Mob at Mt Piere for their permission and co-operation in helping us to
undertake research on their lands; in particular we thank Jimmy Shandley and Lynette Shaw for suggesting and
advice on the usage of the Gooniyandi word ‘girloorloo’. We also thank the staff and managers at Gogo Station for
their time and support, Tracy Sonneman (Western Australia Department of Parks & Wildlife, Broome) for
extensive advice, logistical support and help with permits, and Graeme Armstrong, Paul Horner, Jane Melville,
Tom Parkin, Susan Perkins and Phillip Skipwith for support in the field and photographs. Specimens were
collected and preserved under Western Australian permit numbers SF009528 and SF0009863, and Australian
National University Animal ethics permit number A2013/08. This work was supported by grants from the
Australian Research Council to Paul Oliver, Michael Lee and Paul Doughty, a Melbourne University Collaboration
grant to Jane Melville and Paul Oliver, a McKenzie Postdoctoral fellowship to Paul Oliver from Melbourne
University, an Australian Biological Resources Study grant to Craig Moritz and Scott Keogh and an ARC Laureate
Fellowship to C. Moritz.
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APPENDIX. Comparative (non-type) material examined.
Gehyra girloorloo sp. nov. (n = 6). Western Australia: WAM R175041 (CCM3244), (female), Gogo Station, Limestone Billy
Hills (18.32724°S; 125.76498°E); WAM R175042 (CCM3245), and WAM R175044 (CCM3247), (females), Gogo Station,
Limestone Billy Hills (18.33516°S; 125.75244°E); WAM R175046 (CCM3258) (male), Gogo Station, Pillara Range, south
entrance of Menyous Gap (18.40439°S; 125.83698°E); WAM R175047 (field number—CCM3327), and WAM R175048
(CCM3328) (males), Gogo Station, Virgin Hills, 1 km from road near Bob’s Bore (18.5149°S; 125.9256°E).
Gehyra kimberleyi (n = 19). Western Australia: WAM R108744 (male), 6 km N Gordon Downs Homestead (18.6833°S;
128.5833°E); WAM R108771 (male), Supplejack Bore (18.9166°S; 125.2667°E); WAM R112966 (male), Beagle Bay
Aboriginal Community (17.0730°S; 122.7122°E); WAM R112967 (male), Beagle Bay Aboriginal Community (17.0780°S;
122.7111°E); WAM R112970 (male), Beagle Bay Aboriginal Community (17.0836°S; 122.6597°E); WAM R114212 (female)
and WAMR114213 (male), Cape Leveque (16.4000°S; 122.9333°E); WAM R114370 (female), 17 km SE Gogo Homestead
(18.3333°S; 125.7500°E); WAM R119943 (female), 15 km NE Nifty Mine Site (21.5500°S; 121.6833°E); WAM R154501
(female), Erskine Range (17.8486°S; 124.3425°E); WAM R172737 (male), Windjana Gorge National Park (17.4233°S;
124.9194°E); WAM R172740–172741 and WAM R172744 (females), WAM R172742 and WAM R172743 (males), Windjana
Gorge National Park (17.4233°S; 124.9278°E); WAM R172746 (female), Old Police Station ruins, Windjana Gorge National
Park (17.4258°S; 124.9636°E); WAM R175035 (CCM1332) (male), Meda Station (17.36813°S; 124.00420°E); WAM
R175036 (CCM1336) (male), Birdwood Downs Station (17.34519°S; 123.77831°E).
Gehyra nana (n = 40). Western Australia: NMV D76950 (male), ridge west of Leopold Downs Road (17.90913°S;
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125.28525°E); NMV D76997–76999 (males), Gibb River Road, King Leopold Ranges (17.12977°S; 125.24276°E); NMV
D77006–77007 (females), Gibb River Road, King Leopold Ranges (17.14182°S; 125.23882°E); NMV D77035 (female), NMV
D77036 (male) and NMV D77037–77038 (females), Ngumpan Cliff area (18.75625°S; 126.06474°E); NMV D77042 (male),
The Pinnacles, just off Great Northern Highway (18.73955°S; 125.96368°E); NTM R22918 (female) and NTM R22919 (male),
Spirit Hills, Keep River (15.41666°S; 129.05000°E); NTM R36598 (female), Jasper Gorge, Gregory NP (16.02783°S;
130.78733°E); WAM R108729 (male), Calico Springs, Mabel Downs Station (17.2833°S; 128.1833°E); WAM R108733
(male) and WAM R108735 (female), 11km SE Dave Hill, Mabel Downs Station (17.3000°S; 128.1000°E); WAM R113728
(male), Cadjbut Mine, 90 km SE Fitzroy Crossing (18.7500°S; 126.1500°E); WAM R114449 (female), Koolan Island
(16.1500°S; 123.7500°E); WAM R125993 (female), ~15 km N Mt Septimus (15.5944°S; 129.0186°E); WAM R132858
(female), 10 km S Bow River Station (16.9663°S; 128.2328°E); WAM R140429 (male), ~22 km N Warmun (18.0500°S;
127.6667°E); WAM R172155 (female) Windjana Gorge (17.2272°S; 124.8958°E); WAM R175049 (CCM0534) (male),
Victoria River Escarpment Walk (15.61091°S; 131.11597°E); WAM R175050 (CCM1489) (male), Gibb River Road
(17.13767°S; 125.07829°E); WAM R175051 (CCM1786) (female), Texas Downs Station (17.66931°S; 128.30934°E); WAM
R175052 (CCM1803) (female), Springvale basalt (17.64743°S; 127.69274°E); WAM R175053 (CCM2808) (male), Lake
Argyle rocks (16.11707°S; 128.73813°E); WAM R175054 (CCM2926) (female), East Baines, Gregory NP (15.97138°S;
130.29541°E); WAM R175055 (CCM2984) (male), East Baines camp, Gregory NP (16.45158°S; 130.10257°E); WAM
R175056 (CCM3001) (female), Calcite Flow camp, Gregory NP (16.05033°S; 130.4021°E); WAM R175057 (CCM3033)
(male), WAM R175058 (CCM3039) (male) and WAM R175059 (CCM3040) (male), Sawtooth Gorge (18.42522°S;
127.81967°E); WAM R175060 (CCM3079) (male), Nyulasy (17.76222°S; 129.09862°E); WAM R175061 (CCM3099) (male),
Elephant Rock, Purnululu NP (17.52655°S; 128.36438°E); WAM R175062 (CCM3103) (female), Kurrajong Limestones,
Purnululu NP (17.38542°S; 128.32997°E); WAM R175063 (CMWA78) and WAM R175064 (CMWA79) (females), Springvale
Homestead dump (17.77460°S; 127.69636°E).
Gehyra pilbara (n = 14). Western Australia: WAM R102167 (female), 5.8 km NNW Mt Windell (22.6022°S; 118.5208°E);
WAM R108632 and WAM R108633 (females), 15 km SW Pannawonica (21.8000°S; 116.2333°E); WAM R112678 (male) and
WAM R112705 (female), 7.5 km SSE Onslow (21.7038°S; 115.1278°E); WAM R129420 (female), 1km SE Griffin Gas Plant,
30 km WSW Onslow (21.7833°S; 114.8500°E); WAM R131748 (male), Four Corner's Bore, Hamersley Station (22.3333°S;
117.8667°E); WAM R139441 (male), Mount Minnie (21.3733°S; 115.3733°E); WAM R140311 and WAM R140312 (females),
Millstream Chichester NP (21.4116°S; 117.1561°E); WAM R157285 (female), Yanrey Station (22.2997°S; 114.5931°E); WAM
R158434 (male), Giralia Station (22.6833°S; 114.3667°E); WAM R158438 and WAM R158439 (females), 11 km N Giralia
Homestead (22.5869°S; 114.3872°E).
... The genus has long been recognised as taxonomically difficult (King 1979(King , 1983. Recent phylogeographic studies have revealed extraordinary levels of undescribed cryptic diversity within many nominal species of Australian Gehyra (Sistrom et al. 2013;Moritz et al. 2018;Ashman et al. 2018;Noble et al. 2018;Oliver et al. 2019), and more than 20 new species have been described or elevated from synonymy in the last decade (Sistrom et al. 2009;Doughty et al. 2012Doughty et al. , 2018aDoughty et al. , 2018bHutchinson et al. 2014;Oliver et al. 2016;Bourke et al. 2017;Kealley et al. 2018). As a result of these considerable efforts there are now 42 species of Gehyra recognised in Australia, making it the most speciose genus of gekkotan lizards in Australia, with the rocky Pilbara and Kimberley regions of north-western Australia being centres of diversity and endemism (Ashman et al. 2018;Doughty et al. 2018aDoughty et al. , 2018bKealley et al. 2018). ...
... We recommend the common name 'Amber rock dtella' for this species. Recent taxonomic revisions of Gehyra have moved away from recommending 'dtella' in common names, instead favouring 'Gehyra' (Doughty 2018a;Kealley 2018) or simply 'gecko' (Oliver et al. 2016;Doughty 2018b). Kealley et al. (2018) provide justification for this: "We prefer to use 'Gehyra' as the common name over 'dtella' as the generic name is already available and just as easy or difficult to remember than an additional name fabricated for use as a common name." ...
... nov. is intermediate in size between its two sympatric congeners, G. dubia (larger generalist) and G. einasleighensis (smaller rubble specialist). This is similar to the size and habitat divergence seen between other sympatric Gehyra, such as G. kimberleyi Börner & Schüttler, 1983 (larger generalist), G. girloorloo Oliver, Bourke, Pratt, Doughty & Moritz, 2016 (smaller rock specialist), and G. nana Storr, 1978 (tiny rubble specialists) in the south-western Kimberley region (Oliver et al. 2016;Moritz et al. 2018). This offers further observational evidence that sympatric Gehyra typically differ in body size; however, it is still unclear whether sympatry drives body size divergence, whether those species that have already diverged are more likely to coexist upon secondary contact, or if a combination of these two factors contribute to patterns of body size divergence among Gehyra. ...
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Article
We describe a new species of rock-dwelling Gehyra Gray, 1834 (Gekkonidae) from the Einasleigh Uplands of inland north Queensland, Australia. Morphological, ecological, and molecular data clearly support the new species as distinct and place it within the 'australis group'. Gehyra electrum sp. nov. is distinguished from congeners by a combination of medium adult size (SVL 46-50 mm), an orange-brown to pinkish-orange background colouration with a pattern of distinct whitish spots and irregular black to purple-brown blotches or bars, possessing 7-8 undivided subdigital lamellae on the expanded portion of the fourth toe, and a wedge-shaped mental scale that separates the inner-postmental scales along 40% or more of their length. Gehyra electrum sp. nov. is a rock specialist currently known only from granite outcrops of the Mt Surprise region, Queensland. This is the second recently described Gehyra from the Einasleigh Uplands and adds to the growing number of endemic reptiles recognised in the region.
... King (1979, p. 376) Geckos of the genus Gehyra are among the most taxonomically confusing within the Australian herpetofauna owing to their conservative body form, widely varying characters within taxa, remote locations for many species complexes and poor preservation of colour patterns in museum specimens Sistrom et al. 2009Sistrom et al. , 2014Doughty et al. 2012Doughty et al. , 2018Kealley et al. 2018). Recently, progress has been made resolving the affinities of Gehyra to other gekkonine geckos (Heineke et al. 2011;Gamble et al. 2012;Sistrom et al. 2014) and delimiting species boundaries (Horner 2005;Sistrom et al. 2009Sistrom et al. , 2013Oliver et al. 2010Oliver et al. , 2016Doughty et al. 2012Doughty et al. , 2018Hutchinson et al. 2014). As pointed out 35 years ago, taxonomic progress on Gehyra has been particularly hindered by the loss of colour patterns in preservative and that 'too few specimens from too few localities have been examined' (p. ...
... After this active period in Gehyra systematics, no further species were recognised for over 20 years until the description of G. koira Horner, 2005 (with two subspecies, koira and ipsa) based on morphology. Recent work drawing from combined genetic and morphological data has resulted in many new species descriptions from the Australian Monsoonal Tropics (Doughty et al. 2012Oliver et al. 2016;Bourke et al. 2017) and Central Ranges , and the first proper descriptions of G. lazelli Wells & Wellington, 1983(Sistrom et al. 2009) and G. kimberleyi Börner & Schüttler, 1983. ...
... This integrative approach taken here and elsewhere recently (e.g. Sistrom et al. 2009;Doughty et al. 2012Doughty et al. , 2018Hutchinson et al. 2014;Oliver et al. 2016;Kealley et al. 2018) has proved highly effective at resolving cryptic diversity in conservative Gehyra lineages. ...
... This north-south pattern of species diversity, which appears across multiple groups, has been attributed to an aridity gradient, climate stability or instability, and historical refugia in mesic areas (Afonso Silva et al., 2017;Laver et al., 2018;Palmer et al., 2013). The hypothesis that the presence of rocky refugia drives patterns of diversity is supported by higher diversity along the barrier ranges in the southern Kimberley (Doughty et al., 2018;Oliver et al., 2014Oliver et al., , 2016. ...
... nov.) and one small (D. bennettii), and a smaller generalist woodland species (D. albilabris). Similar body size divergence in sympatric rock-dwelling Gehyra species has been documented in the Kimberley (Moritz et al., 2018;Oliver et al., 2016). In the D. bilineata species group, there is a generalist woodland species (D. margaretae) with smaller body size in the northern Kimberley and a gracile species found in grasslands on river floodplains in the southern Kimberley (D. gracilis sp. ...
... Chromosome evidence from the 1970s and 1980s (King, 1979;Moritz, 1986) indicated several cryptic forms, some of which were described as new species. Beginning with the study of Sistrom et al. (2009), molecular data have unearthed a wealth of phylogenetic diversity in the arid group, while studies in parallel on northern species have also contributed to Gehyra diversity in Australia (Doughty et al., 2012Oliver et al., 2016;Bourke et al., 2017;Moritz et al., 2018). Further studies of Sistrom and colleagues Sistrom, Donnellan & Hutchinson, 2013;Hutchinson et al., 2014) began to clarify the distribution of true G. variegata (Duméril & Bibron, 1836) (type location = Shark Bay in Western Australia (WA)), along with many other cryptic forms, several occurring in the western arid zone and coastal regions. ...
... We tended to not presume species identifications were correct, as the aim was to resolve a cryptic species complex, therefore making prior identifications suspect. We also drew from misidentified specimens that were the focus of other projects (Hutchinson et al., 2014;Oliver et al., 2016). The goal of widespread DNA sequencing was to determine the distribution of species, and to identify putative specimens upon which morphological analysis could be carried out. ...
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Article
The methods used to detect and describe morphologically cryptic species have advanced in recent years, owing to the integrative nature of molecular and morphological techniques required to elucidate them. Here we integrate recent phylogenomic work that sequenced many genes but few individuals, with new data from mtDNA and morphology from hundreds of gecko specimens of the Gehyra variegata group from the Australian arid zone. To better understand morphological and geographical boundaries among cryptic forms, we generated new sequences from 656 Gehyra individuals, largely assigned to G. variegata group members over a wide area in Western Australia, with especially dense sampling in the Pilbara region, and combined them with 566 Gehyra sequences from GenBank, resulting in a dataset of 1,222 specimens. Results indicated the existence of several cryptic species, from new species with diagnostic morphological characters, to cases when there were no useful characters to discriminate among genetically distinctive species. In addition, the cryptic species often showed counter-intuitive distributions, including broad sympatry among some forms and short range endemism in other cases. Two new species were on long branches in the phylogram and restricted to the northern Pilbara region: most records of the moderately sized G. incognita sp. nov. are near the coast with isolated inland records, whereas the small-bodied saxicoline G. unguiculata sp. nov. is only known from a small area in the extreme north of the Pilbara. Three new species were on shorter branches in the phylogram and allied to G. montium . The moderately sized G. crypta sp. nov. occurs in the western and southern Pilbara and extends south through the Murchison region; this species was distinctive genetically, but with wide overlap of characters with its sister species, G. montium . Accordingly, we provide a table of diagnostic nucleotides for this species as well as for all other species treated here. Two small-bodied species occur in isolated coastal regions: G. capensis sp. nov. is restricted to the North West Cape and G. ocellata sp. nov. occurs on Barrow Island and other neighbouring islands. The latter species showed evidence of introgression with the mtDNA of G. crypta sp. nov., possibly due to recent connectivity with the mainland owing to fluctuating sea levels. However, G. ocellata sp. nov. was more closely related to G. capensis sp. nov. in the phylogenomic data and in morphology. Our study illustrates the benefits of combining phylogenomic data with extensive screens of mtDNA to identify large numbers of individuals to the correct cryptic species. This approach was able to provide sufficient samples with which to assess morphological variation. Furthermore, determination of geographic distributions of the new cryptic species should greatly assist with identification in the field, demonstrating the utility of sampling large numbers of specimens across wide areas.
... Gehyra calcitectus sp. nov. is the fourth recently described or redescribed lizard species with a restricted range in the limestone ranges along the southern and eastern fringes of the Kimberley (Oliver et al., , 2016aDoughty, Ellis & Oliver, 2016). Additional limestone endemics lineages from the southern and eastern Kimberley, and the Victoria Rivers District district are likely to represent additional undescribed species (C. ...
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Article
For over two decades, assessments of geographic variation in mtDNA and small numbers of nuclear loci have revealed morphologically similar, but genetically divergent, intraspecific lineages in lizards from around the world. Subsequent morphological analyses often find subtle corresponding diagnostic characters to support the distinctiveness of lineages, but occasionally do not. In recent years it has become increasingly possible to survey geographic variation by sequencing thousands of loci, enabling more rigorous assessment of species boundaries across morphologically similar lineages. Here we take this approach, adding new, geographically extensive SNP data to existing mtDNA and exon capture datasets for the Gehyra australis and G. koira species complexes of gecko from northern Australia. The combination of exon-based phylogenetics with dense spatial sampling of mitochondrial DNA sequencing, SNP-based tests for introgression at lineage boundaries and newly-collected morphological evidence supports the recognition of nine species, six of which are newly described here. Detection of discrete genetic clusters using new SNP data was especially convincing where candidate taxa were continuously sampled across their distributions up to and across geographic boundaries with analyses revealing no admixture. Some species defined herein appear to be truly cryptic, showing little, if any, diagnostic morphological variation. As these SNP-based approaches are progressively applied, and with all due conservatism, we can expect to see a substantial improvement in our ability to delineate and name cryptic species, especially in taxa for which previous approaches have struggled to resolve taxonomic boundaries.
... As it happens the trail of published literature alone supports the taxonomy and nomenclature herein and so I cite it all here. The important published material relevant to the taxonomy and nomenclature of Gehyra sensu lato as defined herein and the decisions made herein are as follows: Andersson (1913), Barbour (1912), Bauer (1994), Bauer and Günther (1991), Beckon (1992), Bobrov and Semenov (2008), Boettger (1895), Bonetti (2002), Schüttler (1982, 1983), Boulenger (1883Boulenger ( , 1885aBoulenger ( , 1885bBoulenger ( , 1887, Brongersma (1930Brongersma ( , 1948, Brown (2014), Brown (1955, Brown et al. (2015), Bourke et al. (2017), Buden and Taboroši (2016), Chan-ard et al. (1999Chan-ard et al. ( , 2015, Chrapliwy et al. (1961), Cogger (2014, Cogger et al. (1983), Crombie and Pregill (1999), Daan and Hillenius (1966), Davies (2012), de Rooij (1915, de Vis (1890), Doody et al. (2015), Doughty et al. (2012), Duméril and Bibron (1836), Duméril and Duméril (1851), Ezaz et al. (2009), Fallend (2007), Fisher (1997), Fitzinger (1843, Flecks et al. (2012), Fry (1914, Garman (1901), Gibbons and Clunie (1984), Girard (1858), Glauert (1955), Goldberg (2014), Gray (1834Gray ( , 1842aGray ( , 1842bGray ( , 1845, Grismer et al. (2007), Günther (1877), Hagey et al. (2017), Hall (2002, Hediger (1933), Heinicke et al. (2011), Horner (2005), Hoser (1989, Hutchinson et al. (2014), King (1979King ( , 1982aKing ( , 1982bKing ( , 1984aKing ( , 1984b, King and Horner (1989), Kinghorn (1924), Kluge (1982Kluge ( , 1993, Kopstein (1926), Laube and Langner (2007), Lesson (1830), Loveridge (1934Loveridge ( , 1948, Low (1979), Lucky and Sarnat (2010), Macleay (1877), Manthey and Grossmann (2007), Maryan (2009), McCoy (2015, Mertens (1974), Meyer (1874), Moritz et al. (2017), Oliver et al. (2010, 2012, 2016a, 2016b, 2017, Mitchell (1965), Oudemans (1894), Peters (1874Peters ( , 1875, Peters and Doria (1878), Pianka (1969), Pianka and Pianka (1976), Ride et al. (1999), Rocha et al. (2009), Rösler (2000, 2017, Rösler et al. (2005), Sang et al. (2009), Shea and Sadlier (1999), Sistrom et al. (2009Sistrom et al. ( , 2012Sistrom et al. ( , 2013, Skipwith and Oliver (2014), Strauch (1887), Steindachner (1867), Sternfeld (1925), Storr (1978Storr ( , 1982, Taylor (1962Taylor ( , 1963, Tiedemann et al. (1994), Tonione et al. (2016), Underwood (1954), Wiegmann (1834), Wellington (1984, 1985), Werner (1901), Wilson and Knowles (1988), Wilson and Swan (2017), Yamashiro and Ota (2005), Zug (1991Zug ( , 2013, Zug and Kaiser (2014), Zug et al. (2011Zug et al. ( , 2012 and sources cited therein. In terms of the nomenclature herein, no names should be altered in any way unless absolutely mandatory under the rules of the International Code of Zoological Nomenclature (Ride et al. 1999). ...
Article
ABSTRACT The lizard genus Gehyra Gray, 1834 as currently recognized consists of roughly 50 recognized species found naturally occurring from mainland south-east Asia to Australia and nearby islands to the north and east including the mid Pacific. This number of currently unrecognized species probably exceeds already described species-level taxa, even though this paper formally names 9 new species and 2 new subspecies, all bar one of which have been confirmed by published molecular data. In spite of the ancient heritage of the assemblage, which is unusual in that numerous species occur on both the Asian and Australian continental plates, divergent lineages with antiquity measured potentially in excess of 25 MYA continue to be treated as being within a single genus. To correct the anomaly, this paper recognizes major divergent species groups as self-contained genera using available and newly created genus names in accordance with the International Code of Zoological Nomenclature (Ride et al. 1999). The assemblage of Gehyra as recognized by most authors to date is herein divided into 14 genera, ten of which are formally named for the first time. The species remaining within Gehyra are further divided into two subgenera, one of which is formally named for the first time. The species within Dactyloperus Fitzinger, 1843 are divided into five subgenera, four of which are formally named for the first time. Another of the newly named genera Edaxcolotes gen. nov. is also divided into two subgenera. All newly named genera and subgenera have divergences of more than 10 MYA from all other species based on numerous published phylogenetic studies. Keywords: Taxonomy; Nomenclature; Lizard; Gekkota; Gekkonidae; Gecko; Dtella; Gehyra; Perodactylus; Peropus; Phryia; Phreodora; Dactyloperus; Asia; Australia; New Guinea; Cambodia; Thailand; new genus; Propemaculosacolotes; Crocodilivoltuscolotes; Edaxcolotes; Extensusdigituscolotes; Brevicaudacolotes; Parvomentumparmacolotes; Papuacolotes; Quattuorunguiscolotes; Colotesmaculosadorsum; Thaigehyra; New subgenus; Halmaherasaurus; Purpuracolotes; Maculocolotes; Wedgedigitcolotes; Saxacolinecolotes; Macrocephalacolotes; species; lacerata, membranacruralis; xenopus; serraticauda; brevipalmata; fehlmanni; oceanica; australis; occidentalis; pilbara; new species; hangayi; paulhorneri; bradmaryani; sadlieri; glennsheai; shireenhoserae; marleneswileae; federicorossignolii; grismeri; new subspecies; bulliardi; graemecampbelli.
... Substantial unrecognised diversity has also been revealed for trees [24], land snails [25], fish [26], frogs [27], and marsupials [28,29]. Meta-analyses of high resolution, comparative phylogeographic diversity across taxa with low dispersal have also revealed new hotspots of endemism that are important to recognise for conservation [30][31][32]. Unrecognised diversity is likely to be especially pronounced for insects in the AMT given that they are so diverse and generally under-studied taxonomically. However, this remains to be investigated. ...
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Article
Ecological opportunity is a powerful driver of evolutionary diversification, and predicts rapid lineage and phenotypic diversification following colonisation of competitor‐free habitats. Alternatively, topographic or environmental heterogeneity could be key to generating and sustaining diversity. We explore these hypotheses in a widespread lineage of Australian lizards: the Gehyra variegata group. This clade occurs across two biomes: the Australian monsoonal tropics (AMT), where it overlaps a separate, larger bodied clade of Gehyra and is largely restricted to rocks; and in the larger Australian arid zone (AAZ) where it has no congeners and occupies trees and rocks. New phylogenomic data and coalescent analyses of AAZ taxa resolve lineages and their relationships and reveal high diversity in the western AAZ (Pilbara region). The AMT and AAZ radiations represent separate radiations with no difference in speciation rates. Most taxa occur on rocks, with small geographic ranges relative to widespread generalist taxa across the vast central AAZ. Rock‐dwelling and generalist taxa differ morphologically, but only the lineage‐poor central AAZ taxa have accelerated evolution. This accords with increasing evidence that lineage and morphological diversity are poorly correlated, and suggests environmental heterogeneity and refugial dynamics have been more important than ecological release in elevating lineage diversity. This article is protected by copyright. All rights reserved
Article
Recent advances in molecular genetic techniques and increased fine scale sampling in the Australian Monsoonal Tropics (AMT) have provided new impetus to reassess species boundaries in the Gehyra nana species complex, a clade of small-bodied, saxicolous geckos which are widely distributed across northern Australia. A recent phylogenomic analysis re-vealed eight deeply divergent lineages that occur as a series of overlapping distributions across the AMT and which, as a whole, are paraphyletic with four previously described species. Several of these lineages currently included in G. nana are phenotypically distinct, while others are highly conservative morphologically. Here we use an integrated approach to ex-plore species delimitation in this complex. We redefine G. nana as a widespread taxon with complex genetic structure across the Kimberley of Western Australia and Top End of the Northern Territory, including a lineage with mtDNA intro-gressed from the larger-bodied G. multiporosa. We describe four new species with more restricted distributions within the G. nana complex. The new species are phylogenetically divergent and morphologically diagnosable, and include the rel-atively cryptic G. paranana sp. nov. from the western Northern Territory, the large-bodied G. pseudopunctata sp. nov. from the southern Kimberley ranges, G. granulum sp. nov., a small-bodied form with granules on the proximal lamellae from the north-west and southern Kimberley ranges and the small-bodied G. pluraporosa sp. nov. restricted to the northern Kimberley. Our revision largely stabilises the taxonomy of the G. nana complex, although further analyses of species lim-its among the remaining mostly parapatric lineages of G. nana sensu stricto are warranted.
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Article
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.
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Article
We describe a new species of large Oedura from the Oscar Range on the southern edge of the Kimberley Craton in north-western Australia. Oedura murrumanu sp. nov. can be distinguished from all congeners by the combination of large size (snout-vent length to 103 mm), moderately long and slightly swollen tail, tiny scales on the dorsum, fringe of laterally expanded lamellae on each digit, and 6-7 paired distal subdigital lamellae on the fourth toe. The new species is the first endemic vertebrate known from the limestone ranges of the southern Kimberley; however, this area remains poorly surveyed and further research (particularly wet season surveys and genetic analyses) is required to better characterise regional biodiversity values.
Article
We describe a new species of velvet gecko (Diplodactylidae: Oedura) from north-east Queensland, Australia. Oedura jowalbinna sp. nov. is a robust, medium-sized (SVL 60–69 mm) gecko that is readily distinguished from its congeners by its distinctive dorsal colour pattern. The dorsum is grey with faint freckling and a pale, dark-edged band across the neck and another across the base of the tail. The combination of a yellow tail and a grey body is also distinctive. Oedura jowalbinna sp. nov. also differs significantly from the most similar congener, O. coggeri, in a multivariate analysis of morphology and scalation, primarily due to its smaller body size, higher interorbital, supralabial and infralabial scale counts, and lower subdigital lamellae scale count. These traits are generally non-overlapping between O. jowalbinna sp. nov. and O. coggeri, however, more individuals of O. jowalbinna sp. nov. need to be assessed to accurately determine variation within the new species. All O. jowalbinna sp. nov. were found at night on overhangs in dissected sandstone escarpment south-west of the town of Laura. Surveys are required to determine the distribution of O. jowalbinna sp. nov. across the sandstone escarpments of the Laura region. This species is the third reptile species (along with the skinks Ctenotus quinkan and C. nullum) described that has a highly localised range centred on the sandstone escarpments of the Laura region. Additionally, included herein is a comparison of O. coggeri and O. monilis. Typical dorsal colour pattern differs between these two species but the large amount of variation (particularly in O. coggeri) merges these differences. Oedura coggeri and O. monilis could not be distinguished in multivariate analyses of morphology and scalation. Genetic data and further analyses of colour pattern, morphology and scalation are required to resolve species boundaries within and between these two species.
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The Australian Brush-tailed Phascogale (Phascogale tapoatafa sensu lato) has a broad but highly fragmented distribution around the periphery of the Australian continent and all populations are under significant ongoing threat to survival. A new appraisal of morphological and molecular diversity within the group reveals that the population in the ‘Top End’ of the Northern Territory is specifically distinct from all others, including those in the Kimberley region of Western Australia to the west and on Cape York of Queensland to the east. The name P. pirata Thomas, 1904 is available for the ‘Top End’ taxon. Three geographically disjunct populations are distinguished at subspecies level within P. tapoatafa on a suite of external and cranio-dental features; these are found in southeast Australia from South Australia to mid-coastal Queensland (nominotypical tapoatafa), southwest Western Australia (wambenger subsp. nov.), and the Kimberley region of Western Australia (kimberleyensis subsp. nov.). A potential fourth subspecies occurs on Cape York but remains too poorly represented in collections for adequate characterization. Molecular divergence estimates based on partial sequences of the mitochondrial cytochrome b gene indicate that the range disjunction across southern Australia probably dates from the Late Pliocene, with the multiple disjunctions across northern Australia being more recent though almost certainly exceeding 400,000 years. An argument is made for the continued use of the subspecies rank in Australian mammalogy, despite a general lack of consistency in its current application.
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The Napier Range of tropical North-Western Australia is an exhumed Devonian reef in an area of intense seasonal rainfall with savanna vegetation. It is composed primarily of three types of hard limestone: a reef facies (the Windjana Limestone), a back-reef facies (the Pillara Limestone), and a fore- and inter-reef facies (the Napier Formation). Slope forms, which range from near-vertical cliffs to gentle convexo-concave profiles, can be related to these facies. Foot-slope pediments are almost ubiquitous. Closed depressions are rare, and fluvial processes are important in shaping the landscape. Surface karren, especially rillenkarren, are well developed on the purer beds. Depositional forms, which include tufas and pseudo-anticlines of calcrete, are notable. Plains fringing the Range show the development of linear aligned gilgai (the orientation of which may be related to prevailing easterly winds) and of degraded Late Pleistocene dunes.
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Species inventories of the main biogeographical areas in the far north of Western Australia are presented. It is shown that the mammal fauna of the Phanerozoic South-west Kimberley reflects the area's intermediate climate, its geographical location between, and its geomorphic continuity with both the sub-humid North Kimberley and the arid Great Sandy Desert. Available data on changes in the mammal fauna of the area since the middle Holocene and since the advent of a pastoral industry in 1890 are presented and discussed.