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RE VI O R A 



Nl ii s e ua in ©i v^omparafive Zoology 



US ISSN 0006-9698 



Cambridge, Mass. 



13 June 2011 



Number 524 



A NEW SPECIES OF THE PACHYDACTYLUS WEBERI COMPLEX 

(REPTILIA: SQUAMATA: GEKKONIDAE) FROM THE 

NAMIBRAND RESERVE, SOUTHERN NAMIBIA 

William R. Branch 1 - 2 , Aaron M. Bauer 3 - 4 , Todd R. Jackman 3 , and Matthew Heinicke 3 

Abstract. A new species of gecko of the Pachydactylus weberi complex is described from the NamibRand 
Reserve in southern Namibia. It is morphologically well differentiated from all other members of this group, in 
lacking thigh tubercles, and can further be distinguished by its small size (< 45 mm snout-vent length), participation 
of the first supralabial in the nostril rim, snout scales much larger than interorbital scales, and three body bands in 
juveniles and adults. It has a distinctive, bold, contrasting hatchling coloration that superficially resembles that of P. 
mclachlani, from which it is distinguished both morphologically and genetically. DNA sequence data from two 
nuclear genes (RAG-1, PDC) and the ND2 mitochondrial gene plus its five flanking tRNAs (2,975 bp) reveal that 
this new species is most closely related, among the taxa sampled, to P. monicae from the lower Orange River valley. 

Key words: Namibia; Gekkonidae; Pachydactylus; new species; ontogenetic color change 



'Port Elizabeth Museum, P.O. Box 13147, Humewood 
6013, South Africa; e-mail: wrbranch@bayworld.co.za 

2 Research Associate, Department of Zoology, Nelson 
Mandela Metropolitan University, P.O. Box 77000, Port 
Elizabeth 6031, South Africa 

3 Department of Biology, Villanova University, 800 
Lancaster Avenue, Villanova, Pennsylvania 19085, U.S.A.; 
e-mail: aaron.bauer@villanova.edu, todd.jackman@ 
villanova.edu, matthew.heinicke@villanova.edu 

4 Research Associate, Department of Herpetology, Mu- 
seum of Comparative Zoology, Harvard University, 26 
Oxford Street, Cambridge, Massachusetts 02138, U.S.A. 



INTRODUCTION 

Geckos constitute a substantial proportion 
of the herpetological diversity of southern 
Africa, with more than 120 species currently 
recorded and at least 16 additional species 
awaiting description. Pachydactylus is the 
most species-rich lizard genus in southern 
Africa, with at least 50 species known from 
the subcontinent (Branch, 1998; Bauer, 
Barts, and Hulbert 2006; Bauer, Lamb, and 
Branch, 2006). Only one species (P. katan- 



The President and Fellows of Harvard College 2011. 



BREVIORA 



No. 524 



ganus of the Democratic Republic of Congo) 
is entirely extralimital to the subcontinent 
(Bauer and Lamb, 2002; Broadley, 2003). At 
least 35 species of Pachydactylus are known 
to occur in the Republic of Namibia, of 
which 18 are endemic (Branch, 1998; Bauer 
et ah, 2002; Bauer, Lamb, and Branch, 
2006). Although most species of Pachydac- 
tylus are morphologically distinct, for many 
years, species boundaries within the small, 
rupicolous species of the P. serval and P. 
weberi complexes were especially problem- 
atic. Relationships within these speciose 
groups were finally teased apart by detailed 
morphological and molecular studies, result- 
ing in the recognition of 22 species (Bauer, 
Barts, and Hulbert, 2006; Bauer, Lamb, and 
Branch, 2006), although one of these, P. 
otaviensis, is now known to belong to 
another clade (Bauer, 2010). 

Despite intensive scientific investigation of 
the Namibian gecko fauna during the last few 
decades, new species remain to be described. 
Shortly after publication of a monographic 
review of the Pachydactylus serval and P. 
weberi complexes (Bauer, Lamb, and Branch, 
2006), a photograph of an unusual hatchling 
gecko was sent to the senior author for 
identification. It was found at Sossusvlei 
Mountain Lodge (now Sossusvlei Desert 
Lodge) in the northern section of the Namib- 
Rand Nature Reserve near the eastern edge 
of the Namib dunefields at the base of the 
Nubib Mountains. It was very similar in color 
pattern to the hatchling of P. mclachlani, a. 
species known only from the Karas Region, 
southern Namibia, approximately 300 km to 
the southeast. This hatchling thus either 
represented another novelty or a considerable 
range extension for P. mclachlani. Subsequent 
visits to the Sossusvlei Desert Lodge resulted 
in the collection of a series of specimens of this 
unusual gecko. Investigation of its morphol- 
ogy and phylogenetic relationships confirm 
that it represents yet another new species of 



the Pachydactylus weberi complex, which we 
describe below. 

MATERIALS AND METHODS 

Morphology. The following measurements 
were taken with Brown and Sharpe Digit-cal 
Plus digital calipers (to the nearest 0.1 mm) 
as per Bauer, Lamb, and Branch (2006): 
snout-vent length (SVL; from tip of snout to 
vent), forearm length (ForeaL; from base of 
palm to elbow), crus length (CrusL; from 
base of heel to knee), tail length (TailL; from 
vent to tip of tail), tail width (TailW; 
measured at base of tail), axilla to groin 
length (TrunkL; measured from posterior 
margin of forelimb insertion to anterior 
margin of hindlimb insertion), head length 
(HeadL; distance retroarticular process of 
the jaw and snout tip), head width (HeadW; 
measured at angle of jaws), head depth 
(HeadD; maximum height of head, from 
occiput to throat), ear length (EarL; longest 
dimension of ear), orbital diameter (OrbD; 
measured at midorbit), nostril to eye distance 
(NarEye; distance between anteriormost 
point of eye and nostril), snout to eye 
distance (SnEye; distance between anterior- 
most point of eye and tip of snout), eye to ear 
distance (EyeEar; distance from anterior 
edge of ear opening to posterior corner of 
eye), and interorbital distance (Interorb; 
shortest distance between left and right 
superciliary scale rows). 

Scale counts and external observations of 
morphology were made with the use of a 
Nikon SMZ-1000 dissecting microscope. 
Comparisons were made with museum ma- 
terial representing all species in the Pachy- 
dactylus serval and P. weberi groups {sensu 
Bauer and Lamb, 2005; Bauer, Lamb, and 
Branch 2006). See Bauer, Lamb, and Branch 
(2006) for a complete list of specimens 
examined. Standard codes for museum col- 
lections follow Leviton et al. (1985). 



2011 



NEW PACHYDACTYLUS FROM NAMIBIA 



Table 1 . Specimens of Pachydactylus used in molecular phylogenetic analyses. All sequences are new to 

this study. 3 



Taxon 



Museum 
Voucher 



GenBank Accession Numbers 



Locality 



ND2 



RAG-1 



PDC 



P. capensis MCZ R- 184499 

P.fasciatus CAS 223931 

MCZ R-185759 



P. serval MCZ R- 185997 

MCZ R-185989 

P. purcelli CAS 198295 

PEM R 16895 

P. mclachlani MCZ R- 186000 
MCZ R- 185094 



P. weberi 



CAS 206824 



PEM R 12449 
P. monicae LSUMZ 57343 

CAS 193418 

P. etultra n. sp. MCZ R- 184977 
MCZ R- 184978 



South Africa: Limpopo, Kgama 
Namibia: Torrabaai Rd., 58 km W of 

Kamanjab (19°39'20"S, 14°21'10"E) 
Namibia: 58 km W of Kamanjab Rest 

Camp on road to Grootberg Pass 

(19°38'57"S, 14°24'33"E) 
Namibia: Brukkaros Mountain, 

S slope (25°53'49"S, 17°46'38"E) 
Namibia: Brukkaros Mountain, 

S slope (25°53'49"S, 17°46'38"E) 
South Africa: Western Cape, Farm 

Oukloof (32°11'36"S, 21°55'38"E) 
South Africa: Northern Cape, Farm 

Kleinbegin (28°54'53"S, 21°40'14"E) 
Namibia: Farm Narudas (27°22'12"S, 

18°51'29"E) 
Namibia: 6.7 km NW of Aroab on 

road to Keetmanshoop (26°46'46"S, 

19°35'17"E) 
South Africa: Northern Cape, 

Brandberg, Farms Kourootje and 

Kap Vley (29°49'52"S, 17°22'35"E) 
South Africa: Northern Cape, 1 .4 km 

S of Garies 
South Africa: Northern Cape, 

Richtersveld National Park, 

Sendelingsdrif 
South Africa: Northern Cape, 

Richtersveld National Park, 

Sendelingsdrif 
Namibia: Sossusvlei Mountain Lodge 

(24°47'2"S, 15°53'22"E) 
Namibia: Sossusvlei Mountain Lodge 

(24°47'2"S, 15°53'22"E) 



HQ 165962 HQ 165992 HQ 165977 
n/a HQ165979 HQ165964 

HQ 165949 HQ 165978 HQ 165963 



HQ165957 HQ165987 HQ165972 

HQ165956 HQ165986 HQ165971 

HQ165955 HQ165985 HQ165970 

HQ 165954 HQ 165984 HQ 165969 

HQ165951 HQ165981 HQ165966 

HQ 165950 HQ 165980 HQ 165965 

HQ165961 HQ165991 HQ165976 

HQ 165960 HQ 165990 HQ 165975 

HQ165953 HQ165983 HQ165968 

HQ 165952 HQ 165982 HQ 165967 

HQ165958 HQ165988 HQ165973 

HQ165959 HQ165989 HQ165974 



Abbreviations for museum numbers are as follows: CAS = California Academy of Sciences, LSUMZ = Louisiana 
State University Museum of Natural Science, MCZ = Museum of Comparative Zoology (Harvard University), PEM 
= Port Elizabeth Museum. 



Molecular Phylogenetics. Sequences were 
obtained from tissue samples representing 
two individuals of the new species, plus two 
each of six other species of the Pachydactylus 
serval and P. weberi complexes (Table 1): P. 
fasciatus, P. mclachlani, P. monicae, P. 
purcelli, P. serval, and P. weberi. Pachydac- 



tylus capensis, a representative of the prob- 
able sister group of this clade (Bauer and 
Lamb, 2005), was chosen as the outgroup. 
The sequence data set includes the complete 
mitochondrial NADH dehydrogenase sub- 
unit 2 gene plus five adjacent tRNAs (ND2; 
1,542 bp aligned), along with portions of the 



BREVIORA 



No. 524 



nuclear protein-coding genes phosducin 
(PDC; 395 bp aligned) and recombination 
activating gene 1 (RAG-1; 1,038 bp aligned). 

Genomic DNA was isolated from ethanol- 
preserved tissue samples with a Qiagen 
DNeasy blood and tissue kit. Polymerase 
chain reaction was then performed on 
isolated DNA in 25-uL reactions with the 
published primers L4437, L4882b, ND2F101, 
ND2R102, H5540, H5934, H6159, PHOF2, 
PHOR1, RAG1F700, RAG1R700, RAG13, 
and RAG18 (Macey et al, 1997, 2000; Groth 
and Barrowclough, 1999; Weisrock et al, 
2001; Greenbaum et ah, 2007; Bauer et al, 
2007). One novel primer was also used: 
ND2F106 (5'-ACAGGATTYATACCAAA- 
ATG-3'). Reaction conditions were as fol- 
lows: denaturation at 95°C (2 min), then 32- 
34 cycles of denaturation at 95°C (35 s), 
annealing at 50°C (35 s), and extension at 
72 °C (initially 150 s, with 4 s added per cycle). 
Annealing temperature was lowered for poor- 
yielding samples. The AMPure magnetic bead 
system (Agencourt Bioscience) was used to 
purify amplified DNA fragments. The se- 
quencing reaction employed dye-terminator 
chemistry with cycle sequencing, followed by 
purification via the CleanSeq magnetic bead 
system (Agencourt Bioscience). An ABI 3700 
sequencer was used for electrophoresis and 
analysis. Resulting chromatograms were in- 
spected by eye and compared against their 
reverse complements in BioEdit (Hall, 1999) 
to detect call errors, with sequences translated 
to check for premature stop codons. Initial 
alignments were performed with Clustal W 
(Thompson et al, 1994) and manually cor- 
rected to take into account codon deletions 
and tRNA secondary structure. The separate 
gene sequences were then concatenated into a 
single data set for analyses. 

Maximum likelihood and parsimony 
methods were used for phylogenetic recon- 
struction. All three gene regions were con- 
catenated for the analyses. Both analyses 



treated gaps as missing data and employed 
pairwise deletion. PhyML (Guindon and 
Gascuel, 2003) and jModelTest 0.1.1 (Posa- 
da, 2008) were used to estimate parameters 
for all standard models of evolution. The 
Akaike Information Criterion (AIC) was 
used to identify the best fitting model of 
evolution (GTR + Y) for use in the likelihood 
analysis. The likelihood analysis was per- 
formed in PhyML version 2.2.4 with a 
BIONJ starting tree, the gamma parameter 
fixed to the previously estimated value, and 
empirical base frequency estimates. The 
parsimony analysis was performed in 
MEGA 4 (Kumar et al, 2008) with the use 
of a min-mini heuristic search (search level 
10). For both analyses, branch support was 
assessed by running 1,000 bootstrap repli- 
cates. 

DESCRIPTION OF NEW SPECIES 

Pachydactylus etultra Branch, Bauer, 
Jackman, and Heinicke, new species 

Sossus Gecko 
Figures 1-3; Table 2 

Holotype. MCZ Rl 84980 (field no. MCZ 
A38619), adult female, Republic of Namibia, 
Hardap Region, Maltahohe District, Namib- 
Rand Nature Reserve, Sossusvlei Desert 
Lodge, 24°46'45.3"S, 15°53'17.8"E (quarter- 
degree unit 2415DD), 896 m elevation, 21 
September 2006, W.R. Branch and P. Dun- 
ning. 

Paratypes. Nine specimens, all from same 
locality as holotype; MCZ R 184977-79, 
184981-82 (four adult females and an adult 
male; see Table 2), 21 September 2006, W.R. 
Branch, A.M. Bauer, J. Marais, & T. Jack- 
man; PEM R 17293 (hatchling), 20 April 
2007, W.R. Branch; PEM R 17294 (sub- 
adult), 21 April 2007, W.R. Branch; PEM 
R 17295 (adult male) and PEM R 17287 
(adult female), 22 April 2007, W.R. Branch, 
D. Branch, & V. Swanepoel. 



2011 



NEW PACHYDACTYLUS FROM NAMIBIA 




Figure 1. Preserved holotype of Pachydactylus etultra, new species (MCZ R 184980). Scale bar = 10 mm. 



Diagnosis. A dorsoventrally flattened spe- 
cies (Figs. 1, 2A) typical of the P. weberi 
complex {sensu Bauer, Lamb, and Branch, 
2006), in having the rostral excluded from 
the nostril (Figs. 2B, C), but distinguished 
from all other species in the complex in 
lacking thigh tubercles (Fig. 2D). It can be 
further distinguished from members of the P. 
weberi complex by a combination of the 
following characters: rupicolous (terrestrial 
in P. fasciatus); the first supralabial narrowly 
enters the nostril (excluded in P. kobosensis) 
(Fig. 2C); snout rounded and nasal region 
moderately inflated (snout pointed and nasal 
region strongly inflated in P. werneri); scales 
on snout larger than interorbital tubercles 
(equal in size in P. monicae) (Fig. 2B); digits 
not long (digits long in P. werneri) (Fig. 2C); 
body slender, caudal tubercles within a tail 
whorl well separated (body large and robust, 
caudal tubercles within a tail whorl abut in 
P. fasciatus); size small, SVL < 45 mm 
(maximum SVL > 49 mm in all other 
members of the complex, except P. robertsi, 
P. reconditus, P. monicae, P. mclachlani, and 
P. visseri); dorsal scalation heterogeneous 
(largely homogeneous in P. kobosensis), with 



the entire dorsum of trunk equally tubercu- 
late (anterior third of dorsum typically less 
strongly tuberculate than posterior in P. 
acuminatus); three body bands (including 
nape and sacral bands) present in juveniles 
(Fig. 3B) and adults, although sometimes 
obscured in the latter (all bands, including 
nape, absent in P. atorquatus; only nape 
band present in juveniles and adults of P. 
robertsi and P. reconditus; four body bands 
in P. werneri; and five or more body bands in 
P. visseri, P. tsodiloensis, and P. water- 
bergensis); bands narrow and straight-edged, 
lost or obscured in adults (body bands broad 
and retained in adults in P. goodi, bands 
broad and wavy-edged in P. weberi). 

Description of Holotype. Adult female. 
SVL 44.5 mm (Figs. 1, 2A, B). Body mod- 
erately depressed, elongate (TrunkL/SVL 
0.41). Head short (HeadL/SVL 0.25), mod- 
erately wide (HeadW/HeadL 0.68), de- 
pressed (HeadD/HeadL 0.37), distinct from 
neck, which is inflated because of enlarged 
endolymphatic sacs. Lores inflated, interor- 
bital region flat. Snout moderately long 
(SnEye/HeadL 0.39), much longer than eye 
diameter (OrbD/SnEye 0.59); scales on snout 



BREVIORA 



No. 524 




Figure 2. Life views of Pachydactylus etultra: adult female holotype (MCZ R 184980) illustrating the depressed 
habitus and characteristic coloration (A) and enlarged scales of the snout and participation of the first supralabial in 
the nostril rim (B). Adult female paratype (PLM R 1 7287) showing the typical somewhat inflated circumnasal scales 
and short digits (C), small dorsal tubercles, and atuberculatc thighs (D) of this species. Photographs by W. R. Branch. 



2011 



NEW PACHYDACTYLUS FROM NAMIBIA 




Figure 3. Life views of Pachydactylus etultra: hatchling paratype (PEM R 17293) showing the characteristic 
boldly contrasting pattern (A). Subadult paratype (PEM R 17294) showing the fading of the dark dorsal background, 
leaving darker margins to the body bands (B). Photographs by W. R. Branch. 



and canthus large, smooth, flattened, het- 
erogeneous in size and shape; scales of 
interorbital and parietal regions strongly 
heterogeneous, with tiny granules inter- 
spersed with larger, domed to conical, 
rounded tubercles, each about one-quarter 
the size of largest snout scales. Enlarged 
conical tubercles regularly scattered across 
occipital and temporal regions. Eye small 
(OrbD/HeadL 0.23); orbits without extra- 



brillar fringes; 4—5 superciliaries at postero- 
dorsal corner of orbit bearing very small 
spines; pupil vertical, with crenellated mar- 
gins. Ear opening oval, horizontal, small 
(EarL/HeadL 0.07); eye to ear distance 
approximately equal to diameter of eye 
(EyeEar/OrbD 1.04). Rostral approximately 
55% as deep (0.6 mm) as wide (1.1), no 
rostral groove, contacted by two enlarged 
supranasals and first supralabials; nostrils 



BREVIORA 



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oval, oriented dorsolaterally, each surround- 
ed by two postnasals, supranasal, and 
narrowly by first supralabial; supranasals in 
broad contact; dorsal postnasal slightly 
larger than ventral postnasal, separated by 
a single granule; nostril rims moderately 
inflated; 1-2 rows of scales separate orbit 
from supralabials; mental elongate with 
nearly parallel sides, 2.25 times deeper 
(1.8 mm) than wide (0.8 mm); no enlarged 
postmentals or chin shields. Enlarged supra- 
labials (left/right) counted to rictus 13/13 (8/8 
to midorbit); infralabials 10/10; interorbital 
scale rows between superciliary rows at 
midpoint of orbit 25 (11 across narrowest 
point of frontal bone). 

Dorsal tubercles large (about six times the 
size of adjacent scales), largest dorsolaterally 
and smallest along dorsal midline and on 
anterior one third of trunk, rounded, with a 
pronounced median keel, forming 14 regular 
longitudinal rows on trunk, grading into 
posteriorly directed, conical scales on lower 
flanks; each tubercle surrounded by a regular 
rosette of about 10 small domed to conical 
scales; ventral scales flattened, rounded to 
oval, subimbricate to imbricate, becoming 
larger posteriorly, largest on posterior abdo- 
men and in precloacal region, approximately 
40 between lowest conical granular rows on 
flank at midbody; tubercular scales on 
dorsum at midbody much larger than ventral 
scales at same level; chin granules approxi- 
mately one fourth size of ventral scales, 
increasing in size rather abruptly between 
gular region and chest. No preanal or 
femoral pores. Scales on palm, sole, and 
ventral surface of forelimb smooth, granular; 
scales on dorsal aspect of forelimb small, 
smooth, heterogeneous, subimbricate; scales 
on preaxial and ventral aspects of thighs 
somewhat enlarged, smooth, imbricate; 
scales on dorsum of thigh nontuberculate, 
heterogeneous, small to large, flattened to 
conical; scales on dorsum of crus small, 



conical, with scattered large (about five times 
the size of adjacent scales), smooth conical 
tubercles. 

Forelimbs short, stout (ForeaL/SVL 0.12); 
hindlimbs short (CrusL/SVL 0.15); digits 
short, minute stylet-like claws visible under 
magnification on all digits except digit I of 
manus; subdigital scansors, except for distal- 
most, entire, present only on distal portion of 
toes, approximately 1.3 times wider than 
more basal (nonscansorial) subdigital scales; 
interdigital webbing absent. Relative length 
of digits (manus): III ~ IV > V > II > I; 
(pes): IV > III > V > II > I. Subdigital 
scansors (excluding small distal divided 
scansor) I (4), II (4), III (4), IV (4), V (4)— 
manus; I (4), II (5), III (5), IV (5), V (5)— pes. 

Tail subcylindrical, weakly depressed; 
original tail somewhat shorter than SVL 
(TailL/SVL 0.91) to somewhat longer than 
SVL for original tails (on the basis of MCZ 
R 184982 and 184978); tail smoothly taper- 
ing, with distinct whorls of scales; dorsal 
caudal scales oval to rectangular, heteroge- 
neous in size, subcaudal scales 3-4 times size 
of dorsal caudals, subimbricate, squarish, 
becoming elongate on distal portion of tail; 
4-5 transverse rows of scales per whorl 
dorsally, 3 ventrally, dorsal scales of poste- 
riormost row of each whorl enlarged, weakly 
keeled, separated from adjacent keeled scales 
by a single, somewhat smaller scale, posteri- 
or half of tail without keels on, or small 
interscales between, enlarged dorsal scales, 
midventral caudal scales enlarged relative to 
adjacent scales (6-8 times size of dorsal 
caudal scales); no postcloacal spurs. 

Coloration of Holotype (in alcohol). Dor- 
sum beige to brownish-yellow with three 
transverse bands bordered by thinner mid- 
brown edges (Fig. 1). Nuchal band extending 
from posteroventral border of orbit, through 
ear, and across nape. Anterior trunk band of 
same width, anterior to midbody, at level of 
elbow of adpressed forelimb. A third, much 



10 



BREVIORA 



No. 524 



broader pale band from anterior to hindlimb 
insertion to anterior sacrum is less conspic- 
uous than the anterior bands, and several 
mid-brown tubercles are enclosed in its pale 
center. Dark anterior border of nape band 
passes through ventral portion of orbit to 
nostril; a diffuse, thick cream stripe from 
anterodorsal margin of each orbit to poste- 
rior margin of ipsilateral nostril. Top of head 
beige with a darker snout and frontal region; 
relatively symmetrical midbrown markings 
at midparietal table and on occiput. Supra- 
labials brown, with areas of reduced pig- 
ment, especially around sutures and on 
posterior scales. Infralabials paler, with 
central regions of some scales deeply speck- 
led, much lighter pigment on others, espe- 
cially mental and first infralabials. Limbs 
mottled, lacking a discrete pattern; digits 
pale. Tail with alternating light (beige) and 
dark (midbrown) bands of approximately 
equal width. Life coloration (Fig. 2A) is 
similar to that in preservative, but the 
background color is a pale lavender. 

Variation. Variation in mensural charac- 
ters of the holotype and adult paratypes are 
presented in Table 2. All paratypes are 
similar in scalation to holotype but have 
more strongly developed keels on the dorsal 
caudal scales. Male paratypes also lack 
precloacal and femoral pores. 

The hatchling paratype (PEM R 17293, 
SVL 17.3 mm) has similar coloration to that 
of hatchling P. mclachlani, with a blackish 
dorsum, wavy-edged bright white trunk 
band, and ashy white nape band that extends 
forward along the jawline to the eye 
(Fig. 3A). As with P. mclachlani juveniles, 
the upper surfaces of the fore- and hindlimbs 
are ashy gray with a brown infusion, whereas 
the top of the head is golden brown. A 
conspicuous pale white eye ring, heavily 
infused with golden orange, is along its 
upper edge. The tail is bright orange with 
8-9 vague darker bands, subequal in width 



to the lighter orange interspaces; it lacks the 
enlarged golden tubercles present in sub- 
adults and adults. Unlike P. mclachlani 
hatchlings, there is an extra ashy white sacral 
band, followed behind by a dark brown band 
at the level of the hindlimb insertion. 
Additionally, there are a number of scattered 
white blotches, including a transverse bar 
almost midway between the chest and sacral 
white bands, and two small white spots on 
either side of the forelimb insertion. An 
additional juvenile (not collected, but pho- 
tographed by Miles Paul) has very similar 
coloration, but the postsacral band is darker, 
with only a single, slightly off-center spot 
between the chest and sacral bands. 

In life, the subadult paratype (PEM 
R 17294) had an intermediate coloration 
between that of the adult and hatchling 
paratypes (Fig. 3B). The body color was a 
pale golden tan with thin dark brown bands 
bordering the dirty cream nape, chest, and 
sacral bands; the enlarged body tubercles 
were golden on the flanks, darker on the 
dorsum. The crown was golden orange with 
darker enlarged tubercles; the extension of 
the nape band onto the snout was white only 
below the eye, and the pale eye ring was 
golden. The upper surfaces of the limbs 
matched the background color of the dor- 
sum, although the limbs were paler at their 
junction with the body; the upper surfaces of 
the digits were grayish, and the ventrum and 
undersurfaces of the head and limbs was 
dirty cream; the tail was diffuse golden 
brown, with little evidence of banding, and 
was highlighted with rings of golden tuber- 
cles. 

Distribution. This species is currently 
known with certainty only from the type 
locality and is therefore another Namibian 
endemic. Suitable habitat occurs extensively 
on rocky mountain slopes of the Nubib 
Mountains in the NamibRand Nature Re- 
serve (Figs. 4, 5), and the species probably 



2011 



NEW PACHYDACTYLUS FROM NAMIBIA 



11 




Figure 4. General habitat at the type locality of 
Pachydactylus etultra. View facing southeast toward the 
Nubib Mountains. Photograph by W. R. Branch. 



has a wider distribution in the region. Several 
specimens from the Maltahohe District 
assigned by Bauer, Lamb, and Branch 
(2006) to P. acuminatus might in fact be 
referable to P. etultra. These include a 
specimen from Sesriem (SAM 44623) and 
two (SAM 45524, 44528) from Tsaris Pass, 
respectively, 35 and 55 km from the type 
locality. A questionable record of P. acumi- 
natus from the Amichab Mountains (but 
without precise locality; see Bauer, Lamb, 
and Branch 2006) in the southern Swakop- 
mund District (TM 32136) could also belong 
to this form and should be re-examined. 

Habitat and Natural History. Some speci- 
mens were collected on rocky slopes at night, 
where they were active on and between rock 
faces. Other specimens were collected around 
the lodge accommodation, some climbing on 
rocks composing the boundary walls and 
chalets. One of the two known hatchlings 
was collected on gravel on a path, whereas 
the other was found on the floor of a chalet, 
both within the lodge complex. Pachydacty- 
lus etultra is the only member of the P. 
weberi species complex found in the region, 
but the more distantly related congeners P. 
rugosus and P. punctatus are expected to be 
present (Branch 1998). 




Figure 5. Map of western southern Africa indicating 
the type locality of Pachydactylus etultra in the northern 
part of the NamibRand Nature Reserve, as well as the 
distribution of all other species of the Pachydactylus 
weberi group. Distributions of other species based on 
Bauer, Barts, and Hulbert (2006) and Bauer, Lamb, and 
Branch (2006). Satellite image from NASA MODIS 
sensor (available at http://visibleearth.nasa.gov). 

Conservation. Several members of the P. 
weberi and P. serval complexes have relative- 
ly restricted distributions, but only one 
species is considered threatened (P. goodi, 
Vulnerable, Bauer, in press). Pachydactylus 
etultra is common at the type locality in a 
well-protected conservancy and with large 
regions of similar habitat in the region. It is 
therefore unlikely to be of conservation 
concern. However, further surveys are re- 



12 



BREVIORA 



No. 524 



100/98 



30/38 



65/32 



86/72 



64/52 



0.05 



■c 



Pachydactylus monicae LSUMZ 57343 
- Pachydactylus monicae CAS 193418 

r Pachydactylus etultra sp. nov. 

I Pachydactylus etultra sp. nov. 
Pachydactylus weberi CAS 206824 
Pachydactylus weberi PEM R12449 
Pachydactylus mclachlani MCZ R-1 86000 



C 



MCZ R-1 84977 
MCZ R-1 84978 



Pachydactylus mclachlani MCZ R-1 85094 
Pachydactylus serval MCZ R-1 85997 
Pachydactylus serval MCZ R-1 85989 
Pachydactylus purcelli CAS 198295 
Pachydactylus purcelli PEM R1 6895 

r Pachydactylus fasciatus CAS 223931 

' Pachydactylus fasciatus MCZ R-1 85759 
Pachydactylus capensis MCZ R-1 84499 



Figure 6. Maximum likelihood phylogeny of Pachydactylus serval and P. weberi group species, rooted with P. 
capensis. Bootstrap values (ML/MP) are indicated at nodes. Support values are not shown for intraspecific 
relationships, but all are 100. 



quired to assess the species' full range and 
habitat requirements, and it should be 
treated as Data Deficient until such infor- 
mation is available. 

Etymology. The specific epithet honors 
"&Beyond" (previously CC Africa), an eco- 
tourism venture that has won many awards 
for the conservation of African wildlife and 
the upliftment of local people. The first 
specimen of this new species was brought to 
our attention by Peter Dunning, then lodge 
manager of the CC Africa Sossusvlei Moun- 
tain Lodge within the NamibRand Private 
Reserve, the area in which all specimens have 
been found. The name is a noun in apposition 
and is a literal Latin translation of "and" (et) 
"beyond" (ultra). 

Phylogenetic Relationships. The final align- 
ment (2,975 bp) included 837 variable and 
635 parsimony-informative sites. A single 
most parsimonious tree was found, with a 
length of 1,721 steps. The best likelihood tree 
had a -In L score of 11,725.40071. Both 
phylogenies displayed identical topologies 
(Fig. 6). Pachydactylus etultra is recovered 
with significant support (maximum likeli- 
hood/maximum parsimony [ML/MP] boot- 



strap 100/98) as a close relative of P. monicae 
from the lower Orange River valley (Fig. 5), 
whereas other between-species branches re- 
ceived nonsignificant support. Both P. mclach- 
lani, the most superficially similar species to 
P. etultra, and P. serval, which shares a similar 
juvenile pattern, fall well outside the etultra + 
monicae grouping. 

DISCUSSION 

Despite more than 50 years of relatively 
intense herpetological study in Namibia (e.g., 
Mertens, 1955, 1971; Bauer et al, 1993; 
Bauer and Branch, 1995; Branch et al, 1996; 
Griffin, 2000, 2003), knowledge of gecko 
diversity in Namibia continues to increase. 
Eleven new species of Pachydactylus have 
been described from Namibia in the first 
decade of this century (Bauer et al, 2002; 
Bauer and Lamb, 2003; Bauer, Lamb, and 
Branch, 2006; Bauer, 2010). This reflects 
greater access to remote areas and richer 
specimen resources on which to base taxo- 
nomic decisions, as well as the availability of 
molecular phylogenetic methods to assess 
relationships. 



2011 



NEW PACHYDACTYLUS FROM NAMIBIA 



13 



The phylogeny presented here is prelimi- 
nary and incomplete, but it does demonstrate 
that P. etultra does not form a monophyletic 
group with the phenetically similar P. mclach- 
lani or with other species with which it shares 
a similar hatchling color pattern. Its closest 
affinity among sampled taxa is with P. 
monicae, a recently described species that 
occurs along the lower Orange River and in 
the valleys of the Fish and Hoolog Rivers. 
The type locality of P. etultra lies approxi- 
mately 360 km NNW of the closest P. 
monicae locality near Rosh Pinah (Bauer, 
Lamb, and Branch, 2006). It is much nearer 
localities for P. acuminatus (not genetically 
sampled here), which occurs from Aus in the 
south to Duwisib (only 85 km from Sossusv- 
lei Desert Lodge) in the north (Fig. 5). A 
more detailed multigene phylogenetic analy- 
sis of the P. serval and P. weberi groups, 
including P. etultra, as well as P. acuminatus, 
P. visseri, P. mclachlani, and P. goodi, all 
taxa not investigated in the cytochrome b- 
based analysis of Bauer, Lamb, and Branch 
(2006), is in preparation. 

The unusual boldly banded black and 
white body coloration of hatchling P. etultra 
is similar to that of a number of other 
Pachydactylus, including P. mclachlani, P. 
carinatus, P. serval, P. bicolor, P. gaiasensis, 
and P. oreophilus. Whereas P. mclachlani is 
relatively closely allied to P. etultra, the other 
species are either members of the P. serval 
complex (P. serval and P. carinatus) or of a 
separate northern Namibian clade (P. bicol- 
or, P. gaiasensis, and P. oreophilus; Bauer 
and Lamb, 2005). Although the species vary 
in size and habitat, they are all mainly 
rupicolous. A similar striking black and 
white-banded hatchling pattern occurs in 
the Bushveld lizard (Heliobolus lugubris), a 
small terrestrial lacertid common in savanna 
habitats, where it is postulated (Huey and 
Pianka, 1977; Schmidt, 2004) that this bold 
hatchling color mimics the coloration of the 



highly distasteful predacious ground beetles 
or "oogpisters" (Anthia sp., Carabidae). 
Although it is possible that a similar form 
of Batesian mimicry could be involved with 
the boldly patterned Pachydactylus hatch- 
lings, protecting them from predation in the 
first few months of life as they disperse from 
their natal oviposition sites, the pattern 
might simply be cryptic and disruptive. 

ACKNOWLEDGMENTS 

We thank Peter Dunning (previously 
Lodge Manager of Sossusvlei Desert Lodge) 
for forwarding Miles Paul's image of the 
unusual hatchling Pachydactylus to WRB for 
identification. We are also grateful to Peter 
and the "&Beyond" staff at the lodge for 
looking after us so well during our fieldwork. 
Vernon Swanepoel and Johan Marais assist- 
ed with the collecting of specimens. We 
thank the collections managers of the Port 
Elizabeth Museum (Gill Watson) and the 
Museum of Comparative Zoology (Jose 
Rosado) for facilitating the study of material 
in their care. Comparative material was also 
made available by Jens Vindum (California 
Academy of Sciences) and Lauretta Mah- 
langu (Ditsong National Museum of Natural 
History [formerly Transvaal Museum], Pre- 
toria). We also thank Mike Griffin, formerly 
of the Ministry of the Environment and 
Tourism, Republic of Namibia, who assisted 
with collecting permits. Specimens were 
collected under Namibian Research/Collect- 
ing Permit 1068/2006. This research was 
funded by grants to AMB and TRJ from 
the National Science Foundation of the 
United States (DEB05 15909 and DEB 
0844523). 

LITERATURE CITED 

Bauer, A. M. 2010. A new species of Pachydactylus 
(Squamata: Gekkonidae) from the Otavi Highlands 
of northern Namibia. Bonn Zoological Bulletin 57: 
257-266. 



14 



BREVIORA 



No. 524 



— . In press. Pachydactylus goodi: species account, 
In M. F. Bates, M. Burger, W. R. Branch, A. M. 
Bauer, J. Marais, G. J. Alexander, and M. S. de 
Villiers eds. Atlas and Red Data Book of the Reptiles 
of South Africa, Lesotho and Swaziland. Pretoria, 
South African National Biodiversity Institute. 

— , and W. R. Branch. 1995. Geographic variation 
in western populations of the Pachydactylus punc- 
tatus complex (Reptilia: Gekkonidae). Tropical 
Zoology 8: 69-84. 

— , and T. Lamb. 2002. Phylogenetic relationships 
among members of the Pachydactylus capensis 
group of southern African geckos. African Zoology 
37: 209-220. 

— , and . 2003. A new species of the 

Pachydactylus weberi group (Reptilia: Squamata: 
Gekkonidae) from the Waterberg Plateau, Namib- 
ia. Cimbebasia 19: 1-12. 

— , and . 2005. Phylogenetic relationships of 

southern African geckos in the Pachydactylus 
Group (Squamata: Gekkonidae). African Journal 
of Herpetology 54: 105-129. 

— , W. R. Branch, and W. D. Haacke. 1993. The 
herpetofauna of the Kamanjab area and adjacent 
Damaraland, Namibia. Madoqua 18: 117-145. 

— , T. Lamb, and W. R. Branch. 2002. A revision 
of Pachydactylus scutatus (Reptilia: Squamata: 
Gekkonidae) with the description of a new species 
from northern Namibia. Proceedings of the Califor- 
nia Academy of Sciences 53: 23-36. 

— , M. Barts, and F. Hulbert. 2006. A new species of 



the Pachydactylus weberi group (Reptila: Squamata: 

Gekkonidae) from the Orange River, with comments 

on its natural history. Salamandra 42: 83-92. 

— , T. Lamb, and W. R. Branch. 2006. A Revision 

of the Pachydactylus serval and P. weberi groups 

(Reptilia: Squamata: Gekkonidae) of Southern 

Africa, with the description of eight new species. 

Proceedings of the California Academy of Sciences 

57: 595-709. 

, A. DE Silva, E. Greenbaum, and T. R. 



Jackman. 2007. A new species of day gecko from 
high elevation in Sri Lanka, with a preliminary 
phylogeny of Sri Lankan Cnemaspis (Reptilia: 
Squamata: Gekkonidae). Mitteilungen aus dem 
Museum fiir Naturkunde in Berlin. Zoologische 
Reihe 83(Supplement): 22-32. 
BRANCH, W. R. 1998. Field Guide to the Snakes and 
Other Reptiles of Southern Africa, 3rd ed. Cape 
Town, Struik Publishers. 

, A. M. BAUER, and D. A. Good. 1996. A review 
of the Namaqua gecko, Pachydactylus namaquensis 
(Reptilia: Gekkonidae) from southern Africa, with 



the description of two new species. South African 
Journal of Zoology 31: 53-69. 

Broadley, D. G. 2003. Pachydactylus katanganus de 
Witte 1953, a species endemic to the Upemba 
National Park (Sauria: Gekkonidae). African Jour- 
nal of Herpetology 52: 69-70. 

Greenbaum, E., A. M. Bauer, T. R. Jackman, M. 
Vences, and F. Glaw. 2007. A phylogeny of the 
enigmatic Madagascan geckos of the genus Uropla- 
tus (Squamata: Gekkonidae). Zootaxa 1493: 41-51. 

Griffin, M. 2000. The species diversity, distribution and 
conservation of Namibian reptiles: a review. Na- 
mibia Wissenschaftliche Gesellschaft Journal 48: 
116-141. 

. 2003. Annotated Checklist and Provisional 

National Conservation Status of Namibian Reptiles. 
Windhoek, Namibia Scientific Society. 

Groth, J. G., and G. F. Barrowclough. 1999. Basal 
divergences in birds and the phylogenetic utility of 
the nuclear RAG-1 gene. Molecular Phylogenetics 
and Evolution 12: 115-123. 

Guindon, S., and O. Gascuel. 2003. A simple, fast, arid 
accurate algorithm to estimate large phylogenies by 
maximum likelihood. Systematic Biology 52: 696- 
704. 

Hall, T. A. 1999. BioEdit: a user-friendly biological 
sequence alignment editor and analysis program for 
Windows 95/98/NT. Nucleic Acids Symposium 
Series 41: 95-98. 

Huey, R. B., and E. R. Pianka. 1977. Natural selection 
for juvenile lizards mimicking noxious beetles. 
Science 195: 201-203. 

Kumar, S., J. Dudley, M. Nei, and K. Tamura. 2008. 
MEGA: a biologist-centric software for evolution- 
ary analysis of DNA and protein sequences. 
Briefings in Bioinformatics 9: 299-306. 

Leviton, A. E., R. H. Gibbs, E. Heal, and C. E. 
Dawson. 1985. Standards in herpetology and 
ichthyology: part 1, standard symbolic codes for 
institutional resource collections in herpetology and 
ichthyology. Copeia 1985: 802-832. 

Macey, J. R., A. Larson, N. B. Ananjeva, Z. Fang, 
and T. J. Papenfuss. 1997. Two novel gene orders 
and the role of light-strand replication in rear- 
rangement of the vertebrate mitochondrial genome. 
Molecular Biology and Evolution 14: 91-104. 
— , J. A. Schulte, A. Larson, N. B. Ananjeva, Y. 
Wang, R. Pethiyagoda, N. Rastegar-Pouyani, 
and T. J. Papenfuss. 2000. Evaluating trans-Tethys 
migration: an example using acrodont lizard 
phylogenetics. Systematic Biology 49: 233-256. 

Mhrthns, R. 1955. Die Amphibien und Reptilien 
Siidwestafrikas, aus den Ergebnissen einer im Jahre 
1952 ausgefiihrten Reise. Abhandlungen der Senck- 



2011 



NEW PACHYDACTYLUS FROM NAMIBIA 



15 



enbergischen naturforschenden Gesellschaft 490: 
1-172. 
— . 1971. Die Herpetofauna Siidwest-Afrikas. Ab- 



handlungen der Senckenbergischen naturforschenden 

Gesellschaft 529: 1-110. 
Posada, D. 2008. jModelTest: phylogenetic model 

averaging. Molecular Biology and Evolution 25: 

1253-1256. 
Schmidt, A. D. 2004. Mimikry zwischen Laufkafern und 

Eidechsen. Frankfurt am Main, Edition Chimaira. 
Thompson, J. D., D. G. Higgins, and T. J. Gibson. 1994. 

CLUSTAL W: improving the sensitivity of pro- 



gressive multiple sequence alignment through se- 
quence weighting, position-specific gap penalties 
and weight matrix choice. Nucleic Acids Research 
22: 4673^680. 
Weisrock, D. W., J. R. MAceY, I. H. Ugurtas, A. 
Larson, and T. J. Papenfuss. 2001. Molecular 
phylogenetics and historical biogeography among 
salamandrids of the "true" salamander clade: rapid 
branching of numerous highly divergent lineages in 
Mertensiella luschani associated with the rise of 
Anatolia. Molecular Phylogenetics and Evolution 18: 
450^164.