Table Of ContentJournal of the Royal Society of Western Australia, 81:201-223, 1998
Morphological and genetic discrimination of new species and subspecies
of gekkonid and scincid lizards (Squamata: Lacertilia)
from the Carnarvon Basin region of Western Australia
K P Aplin1 & M Adams2
1 Department of Terrestrial Vertebrates,Western Australian Museum, Francis St, Perth, WA 6000
email: [email protected]
2 Evolutionary Biology Unit, South Australian Museum, North Terrace, Adelaide, SA 5000 email: [email protected]
Manuscript received February 1998; accepted April 1998
Abstract
Two new species and one new subspecies of lizards are described under the gekkonid genus
Diplodactylus and the scincid genera Ctenotus and Menetia. The gecko Diplodactylus klugei sp nov
and the skink Ctenotus maryani sp nov both qualify as 'cryptic species' in the sense that they are
morphologically similar but genetically distinct from sympatric congeners. The skink Menetia surda
cresswelli subsp nov is morphologically distinct from the typical race but shows only a minimal
level of genetic divergence in allopatry. The three taxa clearly illustrate the frequent lack of
congruence between morphological and genetic differentiation among squamates. A plea is made
for more routine use of allozyme analysis to detect 'cryptic' species within regional vertebrate
faunas.
Introduction examination of voucher specimens obtained from the
Biological Survey sampling quadrats (each with an area
The Carnarvon Basin region of northwest Western
of 16 ha). In the case of the new Ctenotus and
Australia is an area of exceptional reptile diversity with
Diplodactylus species, material provisionally identified as
more than 120 species recorded as a consequence of
C. iapetus Storr and D. pulcher (Steindachner) showed
herpetological surveys conducted during the mid 1970s
subtle but consistent patterns of variation which led to
to early 1980s (Storr & Hanlon 1980; Storr & Harold 1978,
suspicions that the samples might each contain more
1980, 1984, Storr et a\. 1983). In 1995-97 the area was
than one biological species. Material of each potential
resampled as part of the southern Carnarvon Basin
species-composite was initially divided up according to
Biological Survey, a joint project of the Western
quadrat and sex. Starting with one of the more peripheral
Australian Department of Conservation and Land
localities (e.g. most easterly), the material of each sex was
Management and the Western Australian Museum. Close
examined in turn, paying close attention to the nature of
study of the combined regional collections, now totalling
any variation within and between each quadrat sample.
over 11000 specimens, has led to the recognition of new
By working systematically through the voucher material
species within the genera Ramphotyphlops (Typhlopidae),
in this way, it was possible to detect subtle but systematic
Crenadactylus and Diplodactylus (Gekkonidae), Delma
shifts in character states which might otherwise be
(Pygopodidae), and Ctenotus, Cryptoblepharus, Lerista and
overlooked as intrapopulational variation. Each of "C.
Menetia (Scincidae). In most cases, clarification of formal
iapetus" and "D. pulcher" were divided into two putative
taxonomy will require large scale revisionary studies,
species.
several of which are currently in progress. However, in a
few cases the taxonomic issues were largely confined to Allozyme electrophoresis was then used to test for the
the Carnarvon Basin or adjacent regions and thus could presence of more than one biological species in each case.
be resolved without major revisionary work. The 'null hypothesis', that all of the individuals are
drawn from a single, freely interbreeding population,
In this paper we describe two new species and one
would be supported by random allelic variation between
new subspecies of lizards belonging to the genera
the two putative taxa. Rejection of the null hypothesis
Diplodactylus, Ctenotus and Menetia respectively.
(i.e. recognition of discrete, reproductively isolated
Allozyme electrophoresis has been used to demonstrate
populations) would be indicated where significant non-
reproductive isolation between sympatric congenors
random patterns of allelic variation are observed. In the
and/or to determine the level of genetic differentiation
case of sympatric populations, a lack of interbreeding
between allopatric populations.
constitutes strong grounds for recognition of distinct
species, irrespective of which of the various
contemporary species concepts is followed (see Baum
Materials and Methods (1992) for an introduction to this literature; and Frost &
Hillis (1990) for some herptological examples).
Species detection and morphometric analysis
Once the existence of an additional taxon was
The new taxa were initially detected by careful confirmed by allozyme electrophoresis, all available
voucher specimens from the wider study area were
© Royal Society of Western Australia 1998 examined. This phase of the analysis allowed for a wider
201
Journal of the Royal Society of Western Australia, 81(4), December 1998
assessment of the degree of congruence among the Allozyme electrophoresis of liver homogenates was
various 'diagnostic' morphological features and provided conducted on cellulose acetate gels ('Cellogel', MALTA)
an indication of the wider distribution of each taxon. according to the methods of Richardson et al. (1986). The
In the case of Menetia surda, the majority of Carnarvon following enzymes or non-enzymatic proteins exhibited
Basin specimens conformed to a single 'morph', but this zygograms of sufficient activity to proceed for at least
differed from 'typical' M. surda from the northwest of one species-group: aconitate hydratase (Aeon, EC 4.2.1.3),
Western Australia. Specimens from a geographically acid phosphotase (ACP, EC 3.1.3.2), aminoacyclase (Acyc,
intermediate population on Mardathuna appeared EC 3.5.1.14), alcohol dehydrogenase (Adh, EC 1.1.1.1),
morphologically distinct from both northwestern and adenlyate kinase (Ak, EC 2.7.4.3), albumin (Alb),
southern populations. In this case, allozyme carbonate dehydratase (Ca, EC 4.2.1.1), diaphorase (Dia,
EC 1.6.99), enolase (Enol, EC 4.2.1.11), esterase (Esi, EC
electrophoresis was used to estimate the level of genetic
differentiation between the various allopatric (and thus, 3.1.1.?), fructose-bisphosphatase (Fdp, EC 3.1.3.11),
by definition, reproductively isolated) populations. fumarate hydratase (Fum, EC 4.2.1.2), glyceraldehyde-3-
phosphate dehydrogenase (Gapd, EC 1.2.1.12), guanine
The following measurements were taken with vernier deaminase (Gda, EC 3.5.4.3), glutamate dehydrogenase
calipers for all taxa: SVL (snout-vent length); Tail (tail
(Gdh, EC 1.4.1.13), lactoyl-glutathione lyase (Glo, EC
length); and ForelimbL (length of forelimb including 4.4.1.5), aspartate aminotransferase (Got, EC 2.6.1.1),
manus). The hindlimb was measured as HindlimbL
glucose-6-phosphate dehydrogenase (G6pd, EC 1.1.1.49),
(length of hindlimb including pes) for skinks but as
glycerol-3-phosphate dehydrogenase (Gpd, EC 1.1.1.8),
TibialL (length of lower hindlimb, from upper surface of
glucose-6-phosphate isomerase (Gpi, EC 5.3.1.9), alanine
knee to plantar surface of pes) and BifemoralW aminotransferase (Gpt, EC 2.6.1.2), glutathione
(combined upper hindlimb width, measured across the
peroxidase (Gpx, EC 1.11.1.9), glutathione reductase (Gsr,
knees with femora in horizontal plane) for Diplodactylus; EC 1.6.4.2), 3-hydoxybutyrate dehydrogenase (Hbdh, EC
this reflects differences in the most frequent
1,1.1.30), isocitrate dehydrogenase (Idh, EC 1.1.1.42),
preservational posture, namely hindlimbs extended in cytosol aminopeptidase (Lap, EC 3.4.11.1), L-lactate
the skinks but flexed in the gekkos. Two measurments
dehydrogenase (Ldh, EC 1.1.1.27), malate dehydrogenase
were taken of the head in Diplodactylus, namely; HeadL (Mdh, EC 1.1.1.37), 'malic' enzyme (Me, EC 1.1.1.40),
(measured from tip of snout to anterior margin of
mannose-6-phosphate isomerase (Mpi, EC 5.3.1.8),
auricular fossa); and HeadW (maximum width of head).
nucleoside-diphosphate kinase (Ndpk, EC 2.7.4.6), purine-
Except where noted, head scale terminology follows nucleoside phosphorylase (Np, EC 2.4.2.1), dipeptidase
Taylor (1935) for skinks and Kluge (1967) for (PepA, EC 3.4.13), tripeptide aminopeptidase (PepB, EC
Diplodactylus. Paravertebral scales (PV) in skinks were 3.4.11), proline dipeptidase (PepD, EC 3.4.13),
counted from immediately behind the parietal scales phosphoglycerate mutase (Pgam, EC 5.2.4.1),
down to a point opposite the vent. Subdigital lamellae phosphogluconate dehydrogenase (6pgd, EC 1.1.1.44),
(SDL) were counted on the longest finger (3rd; SDLM3) phosphoglycerate kinase (Pgk, EC 2.7.2.3),
and longest toe (4th; SDLP4), from the point of divergence phosphoglucomutase (Pgm, EC 5.4.2.2), pyruvate kinase
of the digit up to, but not including, the claw-sheath. For (Pk, EC 2.7.1.40), superoxide dismutase (Sod, EC 1.15.1.1),
purposes of pattern description of skinks, longitudinal L-iditol dehydrogenase (Sordh, EC 1.1.1.14), and triose-
scale rows are numbered outward from the paravertebral phosphate isomerase (Tpi, EC 5.3.1.1). The nomenclature
series (SR-1) around to the ventral midline (e.g. SR-13). for identifying loci and allozymes is outlined by Adams
Midbody scales rows (MBS) were counted midway et al. (1987).
between the axilla and groin in skinks.
Genetic distances were expressed as percentage fixed
Statistical analyses of individual measurements and differences (% FD, Richardson et al. 1986) and Nei's
scale counts were performed by analysis of variance. unbiased genetic distance (Nei D; Nei 1978).
Before performing interspecific analyses, each sample Heterozygosity values for each taxon were calculated as
was examined for intraspecific sexual dimorphism. In all the proportion of all genotypes observed to be
cases, this was found to be non-significant (i.e. P > 0.05) heterozygotes, averaged over all loci.
for the measured variables. Relative limb lengths were
examined by ratios calculated against SVL; these values
are not directly amenable to statistical analysis (Atchley Systematics
et al. 1976) but provide a useful guide to interspecific
differences in shape. Family Gekkonidae
Unless otherwise indicated, specimen registration Diplodactylus klugei sp nov
numbers refer to the herpetological collection of the
(Fig 1A)
Western Australian Museum. The sex of voucher
specimens was determined by dissection and inspection
Holotype
of gonads.
R120941 in the collection of the Western Australian
Museum, an adult female collected on 11/10/1994 at
Allozyme electrophoresis Biological Survey quadrat W04, 7 km SE of Woodleigh
Outstation in 26° 11' 30" S, 114° 30' 34" E (Fig 2).
Liver samples suitable for allozyme electrophoresis
were removed from freshly-killed lizards in the field or
at the WA Museum. Samples were frozen in liquid Paratypes (Biological Survey quadrats in boldtype)
nitrogen immediately after removal and stored at -80 °C. 14 km W Giralia HS (2-2° 40' S, 114° 14' E): 61089; 26
202
McnBHBuniiinnHBi
K P Aplin & M Adams: Morphological and genetic discrimination of new species and subspecies of lizards
A. Diplodactylus klugei B. Diplodactylus pulcher
Figure 1. Individuals of Diplodactylus klugei sp nov. and D. pulcher (Steindachner), illustrating the close similarity of these probable
sibling species. Details of specimens as follows; D. klugei, WAM R131016, an adult male from 11 km WNW Woodleigh Outstation,
photographed by B Maryan; D. pulcher, an adult female from Kambalda East, photographed by B Maryan. Note the relatively larger
head and eye of D. pulcher.
Figure 2. Distribution of Diplodactylus klugei sp nov and D. pulcher in the Carnarvon Basin area of Western Australia where they occur in
regional sympatry. Diplodactylus pulcher is widely distributed outside of the study area.
203
Journal of the Royal Society of Western Australia, 81(4), December 1998
km W Giralia HS (22^40' S, 114° 07' E): 61265-66; Bullara in Table 1. Males have higher mean values than females
(22° 41' S, 114° 02' E): 60412-13; 2 km E Bullara HS (22° 41' for every dimension but none of the contrasts is
S, 114° 03' E): 61129-32; 3 km W Winning HS (23M0' S, statistically significant.
114° 30' E): 71308; Warroora (23° 29' S, 113° 48' E): 8159-
General head and body scalation finely granular,
60; 5 km SE Gnaraloo HS (23° 51' S, 113° 34' E): 71572;
granules slightly larger on dorsum and forward onto
CU1, 14 km E Cape Cuvier (24°14' 41" S, 113° 32' 07" E):
head. No tubercles or spines present on body or tail.
126905; MR1, 9.8 km E Mardathuna HS (24° 30' 41" S,
Dorsal and lateral surfaces of tail covered in regular rows
114° 38' 13" E): 120715, 123460, 126758; Callagiddy HS
of small triangular scales (apices directed posteriorly).
(25° 03' S, 114° 02' E): 36748, 40669, 41589; MD2, 9.7 km
Ventral surface with smaller, non-aligned rounded
NE Meedo HS (25° 37' 23" S, 114° 41' 39" E): 120982,
scales. Postcloacal sac (housing hemipenes in males)
121013, 122579, 122658, 122666, 124989; MD1, 10.9 km
covered in larger rounded scales; sac relatively smaller in
NE Meedo HS (25° 37' 31" S, 114° 42' 16" E): 121050,
females. Cloacal spine cluster of males consists of 6-10
122527, 122550-51, 122560, 122650, 126691; MD3, 1.25 km
small, conical scales arranged in 2-3 transverse rows;
NE Meedo HS (25° 39' 14" S, 114° 37' 33" E): 121610; 18.4
cluster present in females but scales smaller, rounded
km N Yaringa HS (25° 47' S, 114° 19' E): 71110; 11 km
rather than conical. Preanal region without enlarged
WNW Woodleigh OS (26° 09' S, 114° 26' E): 131016; W04,
scale series or associated pores.
7 km SE Woodleigh OS (26° 11' 30" S, 11411 30' 34" E):
120872, 120884, 120925, 120939, 122961, 125075; W05, Plantar surface of manus and pes finely granular.
13.5 km SE Woodleigh OS (26° 11' 44" S, 114° 25' 24" E): Digits moderately elongate, narrowing at level of
120870, 120881; Overlander Roadhouse (26° 25' S, 114° 28' terminal phalanx, terminating in paired, ovate apical
E): 15278, 55189; 7 km W Hamelin HS (26ll27' S, 114° 08' plates. Fine claws project well past end of apical plates,
E): 54843; 10 M S Shark Bay turnoff. Northwestern visible to naked eye. Undersurface of digits with small,
Highway (26° 32' S, 114° 31' E): 28654; 6 km NE rounded scales in transverse rows, 2-4 per row.
Mungawolagudi (26°47' S, 115°24' E): 60618. Head elongate, narrow and moderately deep. Snout
distinctly 'beaked', weakly grooved between nostrils. Eye
Diagnosis moderately large; supraciliary scales small, triangular
except for 2-3 small conical scales at posterior corner of
A moderate-sized member of subgenus Diplodactylus
eye. Ear aperture small, horizontally ovate.
with rostral and first supralabial both entering nostril;
posterior supralabials not much larger than general head Rostral scale high, rectangular, forming anterior
scales; mental in long contact with first infralabial; digits border of nostril (Fig 3). Well-defined rostral crease
with single pair of enlarged apical plates, claws project extends V - 2/3 way towards lip. First supralabial high,
well forward of plates; other subdigital scales small, forming inferior rim of nostril. Enlarged supranasal scale
rounded, arranged in transverse rows; dorsal pattern forms upper rim of nostril, opposing scales usually in
usually a series of 4-5 large, dark-rimmed pale blotches broad midline contact but occasionally separated by
on red-brown ground colour (blotches are occasionally small internasal scale (e.g. R122961). series of 2-4 small
united into broad, dark-edged vertebral stripe). scales complete posterior rim of nostril. Variable number
of enlarged scales (0-2) located immediately behind
Description supranasals. Second supralabial about V3 height of first.
All successive supralabials much smaller, decreasing in
A slender-bodied, moderately long-limbed terrestrial
size towards oral rictus. Supralabial tally 13-17. First row
Diplodactylus. Tail shorter than head + body, round in
of loreal scales only slightly smaller than supralabials.
cross-section, widening slightly from base and tapering
gradually to tip. Head relatively long and narrow, with Mental elongate, triangular. First infralabial large, in
distinctly 'beaked' snout. SVL to 58 mm; Tail to 33 mm; broad contact with mental and succeeded by 12-13
HeadL to 11.1 mm; HeadW to 9.0 mm; TibialL to 9.0 mm; additional infralabials in decreasing series.
BifemoralW to 20.5 mm. Other statistical data are given Dorsal pattern variable. Most specimens with 4-5
Table 1
Mensural and meristic data for Diplodactylus klugei sp nov and a sample of D. pulcher from the southern Carnarvon Basin region. All
summary values are mean ± standard deviation. Sexual dimorphism was not present in any dimension so data are pooled for both
sexes. Diplodactylus klugei is smaller than D. pulcher in all dimensions except tail length which is approximately equal in the two
species. The ratio data indicate that D. klugei is similar in body shape except for having relatively shorter legs. Diplodactylus klugei has a
significantly higher number of supralabial scales. Values are mean ± standard error with range and/or sample size (n), and with
significance of interspecific difference by ANOVA with F-value and probability (P; NS indicates no significance).
D. klugei sp nov D. pulcher ANOVA
SVL 44.1 ± 6.95 22-55 (n = 49) 50.10 ± 5.81 32-59 (n = 21) F169 = 11.83 (P<0.01)
Tail 30.13 ± 3.23 23-33 (n = 18) 31.25 ± 6.29 22-36 (n = 4) F12] = 0.18 (NS)
HeadL 9.51 ±1.15 6.3-11.3 (n =48) 10.36 ±6.71 8.8-11.4 (n = 19) F]67 = 8.31 (P<0.01)
HeadW 7.43 + 1.03 4.7-9.0 (n = 48) 8.27 ± 0.85 7.1-97 (n = 18) F165 = 9.62 (P < 0.01)
TibialL 7.70 ± 1.00 4.5-9.0 (n = 48) 8.70 ± 0.77 6.2-9.6 (n = 21) F169 = 16.58 (P < 0.001)
BifemoralW 17.91 ±2.01 11.4-20.4 (n = 41) 19.9 ± 2.12 13.9-23.3 (n = 21) F,'62 = 13.46 (P < 0.001)
Tibial / SVL as % 14.2- 19.0 % (n = 7) 15.4-19.4 % (n = 4)
BifemoralW/SVL as % 34.2- 43.9 % (n = 40) 35.9-44.4 % (n = 21)
Supralabials 14.54 ± 1.18 13-17 (n = 24) 13.10 ±1.33 11-16 (n = 19) F142 = 14.07 (P < 0.01)
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K P Aplin & M Adams: Morphological and genetic discrimination of new species and subspecies of lizards
14; supranasals in medial contact. Dorsal pattern a series
of irregular, dark-edged blotches. Flanks marked with
interspersed pale and dark spots. Single yolked follicle
(diameter 7.1 mm) present on each ovary.
Etymology
Named for Professor Arnold Kluge in specific
recognition of his contribution to the systematics of the
genus Diplodactylus.
Distribution
Diplodactylus klugei appears to be confined to the
Carnarvon Basin but has a moderately wide distribution
within this region (Fig 2). It is recorded from
immediately south of Shark Bay north to Giralia and
from near coastal localities inland to Mardathuna, Meedo
and Woodleigh. It is present on Peron Peninsula but has
not been recorded from Edel Land or any of the Shark
Bay Islands. Further north, it is present in near coastal
habitats on the western margin of Lake Macleod.
Genetic discrimination
Allozyme electrophoresis was conducted with liver
samples from five specimens of D. pulcher and seven of
D. klugei. The material selected for analysis comes from
two areas where the two taxa were obtained in regional
sympatry, viz. the Woodleigh and Mardathuna Biological
Survey sites. Nine specimens were screened for 41
presumptive loci, with three others scored only for the 13
most informative loci. The individual allele scores for 20
variable loci are shown in Table 2. The following loci
were invariant within the initial sample of nine
specimens: AcpI, Acp2, Adhl, Akl, Alb, Dia, Gapd, Gda,
G/o, Gotl, Gpi, Gsr, Idhl, Ldh2, Mdhl, Mdh2, Me2, Pgam6,
Pkl, Sod and Tpi.
The two taxa show fixed allelic differences at Aconl,
Figure 3. Rostral scalation of Diplodactylus klugei sp nov (bottom; Acyc, PepA and Pgm2 and display major differences in
WAM R120925) compared with that of D. pulcher (top; WAM
allele frequency at six other loci; Acon2, Est, Gpt, Ihd2, Np
R126707). Note the diagnostic fusion of prenasal and rostral
scales and the absence of an intemasal scale in D. klugei. and Pgml. This equates to a genetic distance of 10 % FD
or a Nei D of 0.258. Even based solely on the four fixed
differences, the odds against the subsamples being
drawn at random from a single, freely interbreeding
large, pale dark-edged blotches on brown to reddish-
population are exceedingly high (the probability of
brown ground colour (80 % of total sample), but
obtaining no heterozygotes at four loci in a sample of 12
occasional specimens with longitudinal anastomoses
animals drawn at random from a "population" in which
between adjacent blotches (8 %; e.g. R120925) or with
each locus has allele frequencies of p = 5/12 and q = 7/12 is
continuous dark-bordered vertebral stripe (12 %; e.g.
((l-2pq)12)4 or 1.3 10'14). Thus the genetic data strongly
R120884). Flanks pale brown to reddish-brown with
support recognition of two distinct biological species
irregular pale spots, occasionally interspersed with dark
within this sample.
spots (e.g. R120941). Tail with 4-5 irregular blotches.
Regrown tails lack blotches, usually have irregular dark Given the broad distributional overlap and high
spotting. Limbs with indistinct pale spotting. Lower degree of morphological similarity between the two taxa,
flanks and venter immaculate. it is of interest to explore the possibility of hybridization
between them. The presence of correlated, fixed allelic
Dorsum of head pale brown to reddish-brown,
differences at four loci makes it quite certain that none of
occasionally with darker central spot (e.g. R120941). Side
the tested individuals represent FI hybrids. However,
of head with broad, pale temporal band which passes
examination of the pattern of heterozygosity between the
through eye to terminate at nostril. Posterior to eye,
two taxa does show some interesting features suggestive
temporal band bordered below by dark brown to black
of possible genetic interaction. In both taxa, the overall
line or diffuse band which terminates below eye.
level of heterozygosity is very similar (Ho= 0.120 ± 0.033
for D. pulcher; Ho = 0.100 ± 0.027 for D. klugei), but the
Details of holotype distribution of heterozygotes at those loci showing major
SVL 51mm; Tail 29 mm; HeadL 10.0 mm; HeadW 7.1 gene frequency differences between the two taxa is
mm; TibialL 8.1 mm; BifemoralW 19.3 mm; supralabials somewhat biased towards D. klugei (Ho = 0.376 ± 0.079 vs
205
Journal of the Royal Society of Western Australia, 81(4), December 1998
hat h Ho = 0.209 ± 0.090 for the loci Acon.2, Est, Gpt, ldh2, Np
ates t Sord ahnedte rozPyggmosl;i tyT daob nleo t 2d)i.f feBr becetawuseee n othvee traaxlla , tlheivs eblisa s oisf
c
di 2 probably not a product of bottle-necking in D. pulcher.
n m
i g ro fl3 R3 (5 n3 Xi Xi X) X) X) X) Instead, it more likely reflects either chance genetic drift
h P
s operating on loci which were polymorphic in the
a
d ml ancestral taxon, strong selection operating on certain loci
A U V u u u
gh. Pg ohentleyr oz(yegitohteers ifno r klhugoemi),o zoyr gao tepsr edion mipnualncthlye r onoer- wfaoyr
ei
dl gk -e . . introgression of pulcher alleles into the klugei genotype.
o P
o This issue cannot be resolved with allelic data and will
W
require analysis of mitochondrial genotypes for any
= d
O Pg further clarification.
W 6
on, D Comparison with other species
Stati Pep Diplodactylus klugei is remarkably similar to D. pulcher
a in general morphology (Fig 1A,B). Indeed, were it not for
n
u A the strong evidence of reproductive isolation provided
ath Pep XXXXX « «3 ns nj ns by the genetic data, the morphological differences might
d
ar well have been ascribed to polymorphism and local
M
= Np phenotypic variation.
A For statistical comparison, the type series of D. klugei
M
was compared with a regionally sympatric sample of D.
2 von Basin survey sites;mined for this locus. Idh2 Ldhl Mpi IXXXX X X fl3 X(13 X(U ’ fl psisrpifnneiihurrg tgleosceannitrhmlo itlaefe lnisraardc aa sbaillla(mi nliTalymst elalc r bynbaessf sllgyrs.eei oimmo omn1Dwnap)si. lha ,plw eeitlTcaronriihhn dttcthie adh s cer atitxfDyhannuclneg .url ra u tsekslh dtygulhepuesipreuogsi rs l enatc ibnhah nsoolaehalittrsysonh ta wir sftli tyutslshhh m resaet(th eFshp rpeiaapogprattr srr reoitda3crsDpi)ta efe;.ol Df nd era.kc atr iblsepnuof y udglnfa lo ercaotifothtl nh elmeiyeearsr
e arer
Tabl Code for Carnpe was not det Got2 Gpt X X i X ! "(2tJ i • rn (Q osnwculraai prtwhrmroa so;wlw araeebindr i daeicnl r or teesnalcmtpaaaitclacievtsa seal w lly( piT tslhlacaa tabretllghesee s er;a 1 fne)ai;ydr se asts ls .ihi sgnAhohfrortltltareyhtlrea o,br lu oingamwehla;e e rrDnll oyt.ann -kclgu loeumsnrgc cbeadieel iaeragl neiiotddnsf
ci. oty D. pulcher both occur in blotched and lined pattern
loen m morphs, the relative frequency of each within the two
ariablethe g Fu tsapxeac imise nvse rbye adri faf esriemnpt.l eI np atDte. rnk luogf epi aoranlllyel 6lo noguitt uodfi n5a1l
v lines, contrasting with 18 out of 23 specimens of the
20 Fdp (1! (5 1 "Jr ¥ ¥ TO regionally sympatric sample of D. pulcher. Four other
at specimens of D. klugei show an intermediate pattern
v
o where the more characteristic blotches are united by a
sp n Est AS fS verterbal stripe.
ei Diplodactylus pulcher has an extensive distribution
g
D. klu Enol (fl (? (? (0 (C (13 (13 oWuetsstiedren Aofu stthrea liCa aerxncaerpvt ofno r Bthaes isno,u tchoevrner ainngd thmeu ncohr tho¬f
d eastern regions (Fig 4). The WA Museum holds a total
n
cher a Acyc X X X X "Jr X X oexf a4m3i0n esdp eacsi mpaernts oof f anD .o npguolcinhge r,s tuwdhyi cohf hgaevoge rabpeheinc
ul variation in this species. In the present context, the most
us p on2 important finding is that D. pulcher consistently has
yl Ae separate prenasal scales throughout its range and
ct
da almost always has one or more internasal scales. In
Diplo Aconl (U (a ra (? (5 X X X X X X X cvoanritarnastst, shthoew sf resiqguneinficcya not fg ebologtrcahpehdic vvsa rilaintieodn , pwatittehr na
of higher frequency of blotched individuals in southern
profiles Site nS Sin Sit)O ^O ^ HQ i^OO^O Otj- Om Oin pteorpmuilnaatilo npsa do fs izDe. apnudlc hreerla. tiLviek elwimisbe , ledniggtiht sl eanrge thsus bajencdt
e n to significant geographic variation in D. pulcher.
Allozym Specimex2e3 ^6u^ O'5oC^cO M5svtNDoC 2O(NvNsc Ii5o(OOfNc)' 5'riO(£Nn- "—6rSS^oiNrt5—snOJ. fi5onN'nO Co5r'OS-o'5loOntNo rO5oftoScXr 5fqI-oNoCt 5tooOnN sDDc.i apllpaoutdilaocchntye lrfu esaa stku luraeg sew,i haisol tlhteho uuosgn hlrye ldioaonbr lsyat hl depi sabtitantesgirusn i soahfne ddr o fslritmorambl
206
K P Aplin & M Adams: Morphological and genetic discrimination of new species and subspecies of lizards
112° 114° 116° 118° 120° 122° 124° 126° 128° 130°
Figure 4. Distribution of Diplodactylus pulcher in Western Australia.
proportions provide a supplementary guide to identity Taxonomic remarks
in the case of Carnarvon Basin specimens.
Cogger et al. (1993) list three taxa as synonyms of D.
Kluge (1967) regarded D. pulcher to be an isolated pulcher. The most important of these in the present
species without close relatives. He included it within his context is Diplodactylus lucasi Fry (a replacement name
vittatus group but noted that it might also have links to for Diplodactylus bilineatus Lucas and Frost). The holotype
the conspicillatus group based on the shared reduction of of Diplodactylus bilineatus Lucas and Frost (MOV D7570)
the labial scales. Both Diplodactylus klugei and D. pulcher is accompanied by a label with the following
show a superficial similarity in body patterning to D. information: Minilya, 80 miles inland from Carnarvon
galeatus Kluge of the Central Australian highlands and Well, 9.5.1903, D.T. Wall / "TYPE" on rear. The specimen
D. steindachneri Boulenger of Eastern Australia, but they is of adult size and appears to be a female based on the
differ from these species in various features including size of the postcloacal spines and sac. According to Kluge
the greater degree of reduction of the labial scales. To (personal communication, 1997), the specimen was in a
date, there has been no comprehensive phylogenetic reasonable condition when he examined it in the mid-
analysis of the genus Diplodactylus and very little 1960s; he reported that the rostral and first supralabial
comparative genetic analysis against which to evaluate scales of MOV D7570 were both excluded from the
the results of the present investigation. nostril (Kluge 1963), a character of D. pulcher as restricted
207
Journal of the Royal Society of Western Australia, 81(4), December 1998
here. Unfortunately, the holotype is now very obtained exclusively on MR1. Similarly on Woodleigh,
dehydrated and brittle and the critical scale boundaries D. pulcher was obtained on quadrat W02, and D. klugei
are no longer visible on the rostrum. As suggested by the on quadrats W04-5. Further, more specific field studies
original specific epithet, MOV D7570 has the strongly are required to determine the extent and basis of any
lined dorsal pattern seen in the majority of Carnarvon habitat partitioning between these morphologically
Basin D. pulcher, but a comparable condition is also similar geckos in the southern Carnarvon Basin.
present in a minority of D. klugei. Features more
During the Biological Survey, Diplodactylus klugei was
reminiscent of D. klugei include an elongate mental in
obtained from the same quadrats as D. squarrosus, D.
extensive contact with the first infralabial and long claws
stenodactylus, D. conspicillatus and D. strophurus.
which protrude beyond the apical plates on all digits (the
Diplodactylus pulcher was found together with this same
latter character might reflect shrinkage of the apical
suite of congenors and additionally with D. alboguttatus.
pads). Minilya is on the eastern side of Lake Macleod,
and both D. pulcher and D. klugei could be expected to
occur in the general vicinity.
Family Scincidae
The apparent admixture of both pulcher and klugei
characters in the holotype of bilineatus is perhaps what Ctenotus maryani sp nov
might be expected of a hybrid individual. If this is indeed
so, D. bilineatus Lucas and Frost would be unavailable (Fig 5A)
under the terms of Article 23h of the International Code Holotype
for Zoological Nomenclature (Anon 1985) which
R126761 in the Western Australian Museum, an adult
disallows individuals of hybrid origin as holotypes of
male collected on 18/3/96 at Biological Survey Quadrat
species-level taxa. However, for the present, we are
MR2, 5.6 km W of Mardathuna Homestead, WA, in 24°
inclined to treat D. lucasi Fry as a junior subjective
26' 36" S 114° 30' 43" E (Fig 6).
synonom of D. pulcher, based principally on Kluge's
record of the rostral scalation at a time when the
Paratypes
specimen was in better condition.
The lectotype of Stenodactylopsis pulcher Steindachner 15 km NE Giralia HS (22° 22' S, 114°29' E): 61185; 3 km
E Giralia HS (22° 41' S, 114" 24' E): 61121-2, 61147; 14 km
(from ' Swan River, WA' = hinterland of Perth) and
holotypes of Diplodactylus pulcher dorsotaeniata Pellegrin N Marrilla HS (22” 51' S, 114° 27' E): 61255; 4 km W
Barradale Roadhouse (22° 51' S, 114° 55' E): 80299; 0.5 km
(from 'Birrigrin, WA' = 50 km NE of Sandstone) and
Diplodactlyus pulcher dorsalis Werner (from Eradu, WA) SE Merlinleigh HS (24° 19' S, 115° 11' E): 99636; MR4, 11.8
km W Mardathuna HS (24° 24' 26" S, 114° 28' 24" E):
have not been examined during this study- However, we
120802, 122777, 126774; MR3, 7.7 km W Mardathuna HS
have examined near-topotypic specimens for each of
(24° 25' 44" S, 114" 29' 60" E): 121555, 122735, 125364;
these forms and believe in each case that they can be
MR2, 5.6 km W Mardathuna HS (24° 26' 36" S, 114° 30'
accomodated within a single, geographically variable
43" E): 120685; KE1, 41 km from Binthalya HS (24° 29'
species. On present evidence, we thus find no grounds to
36" S, 115° 01' 51" E): 120530; 9 km NNW Carey Downs
support Wells & Wellington's (1983, 1985) otherwise
HS (25" 32' S, 115" 26' E): 114313.
unsubstantiated resurrection of D. lucasi and D. dorsalis
from synonymy of D. pulcher.
Diagnosis
Habitat and sympatry A medium-sized Ctenotus with pattern 'simple',
comprising eight to ten pale stripes on a dark-brown to
Diplodactylus klugei clearly occupies a variety of
reddish-brown ground colour; body with 26-28 midbody
different habitats. Specimens collected during the
scale rows; digits of manus and pes with keeled and
Biological Survey were obtained from samphire on the
weakly mucronate subdigital lamellae, 14-17 on manal
edge of Lake Macleod, from the floodplain of the
digit III and 24-26 on pedal digit IV; ear lobules
Wooramel river, from the margin of a claypan on Meedo
triangular, numerous and subequal; dorsal and lateral
and from Acacia grasbyi shrubland on calcareous soils on
pattern terminating on neck, dorsum and temporal
Woodleigh. A common factor among these localities
region of head with indistinct, irregular mottling only;
appears to be the presence of relatively hard, loamy soils
legs with irregular and poorly-developed reticulum.
rather than looser sands. In contrast. Biological Survey
capture records of D. pulcher indicate a preference for
Description
sandier substrates either on dunes or in sandplain
habitats. However, certain of the earlier records of D. A relatively slender-bodied, long-limbed Ctenotus with
klugei come from areas of predominantly sandy rather a predominantly 'simple' pattern of alternating pale and
than loamy substrates (e.g. 5 km SE of Gnaraloo, dark stripes. No sexual dimorphism in body size,
Warroora, Giralia ) and D. pulcher is known to occupy a proportions or colouration. SVL to 54 mm; Tail to 119
wide variety of habitat types across its extensive mm; ForelimbL to 14.9 mm; HindlimbL to 23.9 mm.
geographic range.
Paravertebral scales 51-62, mean. = 55.8 ± 3.3 (sd);
Although D. klugei and D. pulcher are regionally scales in 26-29 rows at midbody, modal count 28 (41% of
sympatric over quite a large area, the two species were sample). Other mensural and meristic data are given in
not found to co-occur on any of the Biological Survey Table 3.
quadrats. On Mardathuna, for example, D. pulcher was
Plantar surface of manus and pes spinose,
captured on quadrats MR2-5 while D. klugei was
unpigmented. Digits slender, relatively elongate.
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K P Aplin & M Adams: Morphological and genetic discrimination of new species and subspecies of lizards
A. Ctenotus maryani B. Ctenotus iapetus
Figure 5. Individuals of Ctenotus maryani sp nov. and C. iapetus Storr, illustrating the remarkable similarity of these genetically quite
distinct species. Details of specimens as follows; Ctenotus maryani sp nov, WAM R114313, an adult male from 9 km NNW Carey
Downs Homestead, photographed by B Maryan; C. iapetus, WAM R51018, an adult female from 3 km NW Bullara Homestead,
photographed by R E Johnstone.
112° 113° 114° 115° 116° 112° 113° 114° 115° 116°
C. maryani
Figure 6. Distribution of Ctenotus maryani sp nov and C. iapetus in the Carnarvon Basin region of Western Australia. Both species are
endemic to the region.
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Journal of the Royal Society of Western Australia, 81(4), December 1998
Table 3
Mensural and meristic data for Ctenotus maryani sp nov and C. iapetus. All values are mean ± standard deviation, followed by range
and sample size. Sexual dimorphism was not present in any dimension so data are pooled for both sexes. Ctenotus iapetus evidently
attains a larger overall size than C. maryani but sample means for SVL are not significantly different. Ctenotus maryani has a significantly
higher mean number of paravertebral scales than does C. iapetus. The ratio data indicate that Ctenotus maryani has relatively shorter
limbs than C. iapetus. Values are mean ± standard error with range and/or sample size (n), and with significance of interspecific
difference by ANOVA with F-value and probability (P; NS indicates no significance).
C. maryani C. iapetus ANOVA
SVL 47.56 ± 6.55 33-54 (n = 18) 49.20 ± 11.76 31-68 (n = 82) F1<n = 0.326 (NS)
Tail 98.80 ± 19.98 64-119 (n = 10) 95.84 ± 24.61 60-149 (n = 44) F' = 1.341 (NS)
ForelimbL 12.62 ± 1.77 8.1-14.9 (n = 18) 13.89 ± 2.59 8.9-18.1 (n = 81) F]98 = 3.88 (NS)
PlindlimbL 21.31 ± 2.43 14.8-23.9 (n = 18) 23.23 ± 4.47 15.5-31.0 (n = 77) F]94 = 3.11 (NS)
Tail/SVL * 100 179-236 % 163-244 %
ForelimbL/SVL*100 22.5-29.4 % 22.3-37.9 %
HindlimbL/SVL*100 40.4-52.1 % 39.9-60.6 %
PV 55.83 ± 3.31 51-62 (n = 18) 50.91 ± 2.90 44-61 (n = 70) F = 38.79( P < 0.001)
SDL-M3 16.0 ± 0.71 15-17 (n = 9) 15.66 ± 1.22 13-18 (n = 77) F = 0.668 (NS)
SDL-P4 25.22 ± 1.31 21-26 (n = 18) 26.17 ± 1.63 23-30 (n = 77) Fim = 5.288 (NS)
terminating in fine claw. Subdigital lamellae keeled and interparietal. Parietal eye clearly visible at rear of
with small mucrons. SDL 14-17 on manal digit III, modal interparietal. Parietals in contact behind interparietal.
count 16 (56 % of sample); 24-26 on pedal digit IV, modal Nuchals in 2-4 pairs, 83 % with 4 nuchals on at least one
count 26 (61 % of sample). side.
Ear aperture moderately large. Usually with 2-3 large
Nasals and prefrontals both in broad medial contact.
Loreals 2, anterior higher but less elongate than second triangular lobules, bordered above and below by one or
(Fig 7). Supralabials 8-9, 5 or 6 of these making up more smaller, rounded lobules (Fig 7).
discrete preorbital series. Postsupralabials [postlabials of Ground colour of body dark-brown on dorsum, paler
Taylor (1935)] 2. Infralabials 8. Supraoculars 4, second reddish-brown on flanks. Dorsum with four continuous
wider than third, all but last in contact with frontal (Fig stripes (Fig 9); white paravertebral stripe (positioned
8). Supraciliaries 7-8, either l>2>3»4-7 with anterior 3 centrally on SR-1), off-white dorsal stripe (centre SR-2),
in contact with first supraocular, or l>2>>3-8 with white dorsolateral (centre SR-3) and off-white upper
anterior 4 in contact with first suprocular. Eyelid mobile, lateral (lateral SR-4). Flank always with broad midlateral
with three enlarged scales forming a divided window. and lower lateral stripes; both are white, positioned on
Preoculars 2 [lower is anterior presubocular of Taylor scale rows 6 + part 7 and 8 + part 9 respectively. Lower
(1935)]. Single presubocular, wedged between lateral stripe faintly delineated below by thin brown line
supralabials 5-6 or 6-7. Postsuboculars 3, abutting (centre SR-9). Upper lateral zone variable: either solid
penultimate supralabial and primary temporal, in series brown, covering whole of SR-5 and intercalcating tips of
with pretemporals. Single primary temporal, wedged rows 4 and 6 (e.g. R80299, R121555) or divided by off-
between last two supralabials. Secondary temporals 2, white accessory upper lateral stripe (centre SR-5).
upper much larger than lower. Frontoparietal larger than Accessory stripe usually indistict, comprised of irregular
Figure 7. Scalation of the lateral surface of the head of Ctenotus maryani sp nov (WAM R126761).
210
