Review of the taxonomy of the spiny-tailed lizards of

Transcription

FAUNA OF ARABIA 23: 435–468
Date of publication: 15.07.2007
Review of the taxonomy of the
spiny-tailed lizards of Arabia
(Reptilia: Agamidae: Leiolepidinae: Uromastyx)
Thomas M. Wilms and Wolfgang Böhme
A b s t r a c t : Currently six species of the genus Uromastyx (Reptilia: Agamidae: Leiolepidinae), representing three phylogenetic
lineages, are known to occur in Arabia: Uromastyx aegyptia, U. benti, U. leptieni, U. ornata, U. thomasi and U. yemenensis. The
present paper gives an overview of the taxonomy of these lizards and presents new data on the morphology and ecology of Uromastyx leptieni.
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(Leiolepidinae: Uromastyx :�� :��)
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(Leiolepidinae :�� :��) Uromastyx �� :��
.U. yemenensis � U. benti, U. leptieni, U. ornata, U. thomasi, Uromastyx aegyptia ��
.Uromastyx leptieni ��
INTRODUCTION
Spiny-tailed agamas are small to medium-sized, ground- or rock-dwelling lizards. Most species
reach a maximum length of 25‑50 cm, and only species of the Uromastyx aegyptia group can reach
a total length of up to 70 cm or more. The animals have a bulky, depressed body and strong, short
limbs. The tail is covered by spiny scales, arranged in distinct whorls. The tympanum is visible.
None of the species has a nuchal or dorsal crest or a gular pouch or fan. Only a transverse fold at
the throat is present (gular fold). Body scales are small and mostly homogenous, but some species
have enlarged tubercular scales on the body and/or limbs.
The main diagnostic character of the genus is the highly specialised tooth-like bony structure
replacing the incisor teeth in the upper jaw in adults. This transformation of the premaxillary bone
to a tooth-like structure is an autapomorphy of the genus Uromastyx, convergent in Sphenodon,
which has also a convergently acrodont dentition. Owing to morphological, ethological and geAuthors’ addresses:
Thomas M. Wilms, Reptilium – Terrarien- und WüstenZoo, Werner-Heisenberg-Str. 1, 76829 Landau, Germany; e‑mail: thomasmwilms@
yahoo.de
Wolfgang Böhme, Zoologisches Forschungsmuseum A. Koenig, Adenauerallee 160, 53113 Bonn, Germany; e‑mail: W.Boehme.
ZFMK@uni-bonn.de
436
T.M. Wilms & W. Böhme
netic synapomorphies, butterfly agamas of the genus Leiolepis are considered to be the closest living
relatives of spiny-tailed lizards and both genera are therefore placed in the subfamily Leiolepidinae
(synonym: Uromastycinae) (Peters 1971, Moody 1980, Böhme 1988, Schmitz et al. 2001, Amer
& Kumazawa 2005).
Lizards of the genus Uromastyx occur in the desert belt of the Old World between 5°N and
35°N. Their range covers an enormous land mass, including northern Africa, the Middle East,
Arabia, Iran, Iraq, Afghanistan, Pakistan and north-western India (Fig. 1). Today, 17 species are
recognised, of which six species (with three additional subspecies) are native to Arabia (Kingdom
of Saudi Arabia, Yemen, Sultanate of Oman, United Arab Emirates, Qatar and Kuwait (Wilms
2001, 2005; Wilms & Schmitz 2007).
The species of spiny-tailed lizards currently known to occur in Arabia are: Uromastyx aegyptia
(Forskål, 1775), U. leptieni Wilms & Böhme, 2000, U. benti (Anderson, 1894), U. ornata von
Heyden, 1827, U. yemenensis Wilms & Schmitz, 2007 and U. thomasi Parker, 1930. They represent three different phylogenetic groups, with species compositions listed below (Wilms 2001,
Wilms & Schmitz 2007).
Uromastyx habitats are generally characterised by high temperatures, low precipitation, sparse
vegetation and marked seasonal changes (Plates 1‑2). However, in the coastal regions of south and
south-western Arabia (Asir and Hejaz Mountains, Hadramaut and Dhofar), Uromastyx habitats are
more mesic because of the influence of the summer monsoon, resulting in high air moisture and
elevated precipitation. These habitats are often well vegetated. Uromastyx species are mostly found
on firm soil or on rock surfaces that provide shelter in the form of cracks and crevices. They also
need, at least for part of the year, a sufficient amount of vegetation as a food source. Uromastyx
live mainly in rocky desert and semi-desert habitats that are covered with rocks, scattered stones
or gravel. Large continuous dune areas (erg) with drifting sand do not offer suitable habitats and
represent range barriers. However, this does not prevent Uromastyx from travelling a few kilometres
across sand from one gravel plain to another. Most species are digging, ground-dwelling animals,
while some species live in rocky or mountainous areas and retreat into cracks and crevices. Some
species have even been observed climbing acacia trees (U. aegyptia, U. dispar, U. ocellata and U. ornata). Uromastyx are opportunistic herbivores, which means, that they feed mainly on plants and
to a lesser extent on animals. By far the largest part of the diet is leaves, flowers, sprouts and seeds
of various plant species. Spiny-tailed agamas are strictly diurnal.
MATERIALS AND METHODS
Morphological sampling
The taxonomy presented in the present paper is based on a revision of the genus Uromastyx (Wilms
1998; Wilms & Böhme 2000 a, 2000 b, 2000 c; Wilms & Schmitz 2007), in which a total of
625 specimens have been examined, including 180 specimens of Arabian species. The specimens
are kept in museum collections indicated by the following abbreviations:
BMNH
British Museum of Natural History, London, UK (now: The Natural History
Museum, London, UK, NHM)
MHNG Muséum d’histoire naturelle de la Ville de Genève, Geneva, Switzerland
MNHNP Muséum national d’Histoire naturelle, Paris, France
MTD
Staatliches Museum für Tierkunde, Dresden, Germany
Review of the taxonomy of the spiny-tailed lizards of Arabia
437
Fig. 1: Distribution range of the genus Uromastyx (after Wilms 2001).
MZUF
NHMW
SMF
ZFMK
ZMB
ZMH
ZSM
Università di Firenze, Museo Zoologico “La Specola”, Florence, Italy
Naturhistorisches Museum Wien, Vienna, Austria
Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt a.M., Germany
Zoologisches Forschungsmuseum Koenig, Bonn, Germany
Museum für Naturkunde der Humboldt-Universität zu Berlin, Germany
Zoologisches Institut und Zoologisches Museum der Universität Hamburg, Germany
Zoologische Staatssammlung München, Munich, Germany
Lists of examined specimens have been published by Wilms & Böhme (2000 a, 2000 b, 2000 c),
Wilms et al. (2002) and Wilms & Schmitz (2007).
To obtain new information on the morphology and ecology of Uromastyx leptieni, data were
collected by Thomas Wilms and Brien Holmes at a site near Abu Dhabi International Airport
(United Arab Emirates) between 11 June 2005 and 16 July 2005, including morphometric and
meristic data of 82 live specimens. These specimens were subsequently released in the course of a
translocation project, during which a total of 213 specimens were translocated (Tourenq 2005,
C. Drew pers. comm.).
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Plate 1: Habitat of
Uromastyx thomasi
at Ras Hilf, Masirah
Island, Oman. Photo:
T. Wilms.
Plate 2: Habitat of
Uromastyx aegyptia
microlepis in Mahazat
as-Sayd, Saudi Arabia.
Photo: F. Krupp.
Characters
For each museum specimen the following data were routinely recorded:
– snout-vent-length (SVL)
– tail length
– head width, measured between the anterior margins of the ear openings
– head length, measured from the tip of the snout to the anterior margin of the ear opening
on the left side
– tail width between the 4th and 5th whorl
– maximum tail width at the 5th whorl
– number of scales around mid-body
– number of scales between gular and inguinal fold (= ventrals)
– number of gular scales (from the mental scale to a line between the anterior margins of the
ear openings)
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– number of scales from the mid of the lower end of the ear opening to the mental scale (left
and right)
– number of scales between supralabial and enlarged subocular scale
– number of scales around the 5th whorl
– number of tail whorls
– number of scales beneath the 4th toe on the left side
– number of preanofemoral pores on either side
– number of enlarged scales at the anterior margin of the ear opening
– number of scales from the upper to the lower end of the left ear opening (approximately
three scale rows before the anterior margin of the ear opening)
– number of scales from the upper end of the left ear opening to the first enlarged subocular
scale
– intercalary scales between the whorls (presence or absence)
– presence or absence of enlarged tubercular scales at the flanks and on the dorsum
With the exception of the number of scales around the mid-body, the same measurements and
scale counts as in museums specimens were taken from the 82 live Uromastyx leptieni.
Statistical analyses
The statistical packages Excel 2000, SPSS (10.0) and R 2.2.1 (R Development Core Team 2005)
were used to run the analyses. Two multivariate procedures were selected to evaluate data and to
explore relationships within the Uromastyx aegyptia group (sensu Wilms & Böhme 2000 a).
Hierarchical cluster analysis
Hierarchical cluster algorithms find successive clusters using previously established clusters and
thus grouping similar objects together. Because of the many different methods to calculate distances in a given matrix and the large number of clustering algorithms this method is much more
subjective than the ordination techniques such as PCA (Rastegar-Pouyani 2005). In the present
study the use of average linkage method has produced the highest cophenetic correlation coefficients and the highest γ‑coefficients. Both coefficients provide a measure for the reliability of a
cluster analysis and, according to Handl (2002), a γ‑coefficient larger than 0.70 indicates that the
fit of the analysis is satisfactory or better. A cophenetic correlation coefficient above 0.750 is considered good fit (Rastegar-Pouyani 2005).
Principal component analysis (PCA)
Principal component analysis (PCA) is a multivariate procedure which rotates the data such that maximum variabilities are projected onto the axes. Essentially, a set of correlated variables are transformed
into a set of uncorrelated variables which are ordered by reducing variability. The uncorrelated variables
are linear combinations of the original variables, and the last of these variables can be removed with minimum loss of real data. The main use of PCA is to reduce the dimensionality of a dataset while retaining
as much information as is possible. Results of PCA do not depend on prior specimen classification.
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SYSTEMATIC ACCOUNT
Genus Uromastyx Merrem, 1820
Within the genus Uromastyx, six different species groups have been identified, three of which occur
in Arabia (Wilms 1998; for a proposed phylogeny of the Arabian taxa see Fig. 2):
Uromastyx hardwickii group
Uromastyx asmussi group
Uromastyx princeps group
Uromastyx ocellata group
Uromastyx aegyptia group
Uromastyx acanthinura group
The Arabian representatives of three of these groups (U. princeps group, U. ocellata group and
U. aegyptia group) can be identified by means of the following key.
Key to the Arabian species of the genus Uromastyx
1
–
Preanal and femoral pores absent............................................................................... Preanal and femoral pores present.............................................................................. 2
4
2
–
143‑187 scales around mid-body (average 160.05 ± 8.98), 66‑86 ventral scales
Uromastyx benti
163‑227 scales around mid-body (average 192.53 ± 16.63), 79‑97 ventral scales....... 3
3
Ground colour light brown with five distinct cross-bands on the back (see Plate 15)
Uromastyx yemenensis yemenensis
Ground colour dark brown with light brown dots tending to form transverse rows on
the back (see Plate 17)
Uromastyx yemenensis shobraki
–
4
–
Tail short, up to 35 % of snout-vent-length, disc-shaped
Uromastyx thomasi
Tail long, more than 60 % of snout-vent-length, 22‑27 whorls.................................. 5
5
–
247‑322 scales around mid-body............................................................................... 138‑193 scales around mid-body............................................................................... 6
7
6
110‑150 ventral scales, preserved juveniles reddish brown with a dark vermiculation
or with small irregularly arranged dark and light spots
Uromastyx leptieni
126‑193 ventral scales, preserved juveniles greyish brown with yellowish dots on the
back arranged in transverse rows................................................................................ 8
–
7
–
th
Ratio of tail length to maximum tail width at the 5 whorl: 3.61‑5.30
Uromastyx ornata ornata
Ratio of tail length to maximum tail width at the 5th whorl: 3.03‑3.96
Uromastyx ornata philbyi
8
126‑158 ventral scales, enlarged lateral tubercles present
Uromastyx aegyptia aegyptia
–
149‑193 ventral scales, enlarged lateral tubercles absent
Uromastyx aegyptia microlepis
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Fig. 2: Proposed phylogeny of the Arabian
taxa of the genus Uromastyx based on morphological characters.
Within the Uromastyx aegyptia group, four different taxa have been described: the recently discovered U. occidentalis Mateo, Geniez, López-Jurado & Bons, 1998, from Western Sahara and U. leptieni Wilms & Böhme, 2000 from Oman and the United Arab Emirates and the two well-known
taxa U. aegyptia Forskål, 1775, and U. microlepis Blanford, 1874. Wilms & Böhme (2000 a)
provide a detailed discussion of the taxonomic history of the taxa of the Uromastyx aegyptia group
and recognise three taxa as valid species (U. aegyptia, U. leptieni and U. occidentalis). The fourth
nominal taxon, microlepis, has been treated as a subspecies of U. aegyptia.
The main characters to distinguish the members of the Uromastyx aegyptia group from other
Uromastyx species groups are their large body size (exceeding 700 mm in adult specimens), combined with extraordinarily small body scales.
Uromastyx aegyptia (Forskål, 1775) – Egyptian spiny-tailed lizard
Lacerta aegyptia Forskål, 1775. — Descriptiones animalium avium, amphibiorum, piscium, insectorum, vermium; quae in
itinere orientali observavit Petrus Forskål: 13.
Two subspecies are recognised: Uromastyx aegyptia aegyptia and Uromastyx aegyptia microlepis.
Uromastyx aegyptia aegyptia (Forskål, 1775)
Plate 3
Lacerta aegyptia Forskål, 1775. — Descriptiones animalium avium, amphibiorum, piscium, insectorum, vermium; quae in
itinere orientali observavit Petrus Forskål: 13.
Lacerta harbai Forskål, 1775. — Descriptiones animalium avium, amphibiorum, piscium, insectorum, vermium; quae in
itinere orientali observavit Petrus Forskål: 9 (?; synonymy fide Merrem 1820).
Stellio spinipes Daudin, 1802. — Histoire naturelle, générale et particulière des reptiles. Vol. 4: 31.
Uromastyx spinipes. — Merrem 1820; Versuch eines Systems der Amphibien – Tentamen systematis amphibiorum: 56
(n. comb.).
Lacerta herbai. — Merrem 1820; Versuch eines Systems der Amphibien – Tentamen systematis amphibiorum: 56 (nomen
substitutum pro Lacerta harbai Forskål, 1775).
Mastigura spinipes. — Fleming 1822; The Philosophy of Zoology; or, A General View of the Structure, Functions, and Classification of Animals. Vol. 2: 277 (n. comb.).
Uromastix spinipes. — Boulenger 1885; Catalogue of the Lizards in the British Museum (Nat. Hist.). Vol. 1: 407.
Uromastix aegyptius. — Anderson 1896; A Contribution to the Herpetology of Arabia, with a Preliminary List of the Reptiles
and Batrachians of Egypt: 79, 85.
Uromastyx aegyptica. — Khalil & Hussein 1962; Bulletin of the Zoological Society of Egypt 17: 80.
442
Ty p e s p e c i m e n : Neotype, adult , Egypt, Suez at the road to Cairo, VIII.1982, I. Rehak, ZFMK 44216, designated by Wilms & Böhme (2000 a).
D e s c r i p t i o n : S i z e : Large-growing taxon, maximum total length exceeding 700 mm.
S c a l a t i o n : 247‑322 scales around mid-body, 126‑158 scales between gular and inguinal
fold, 33‑59 gular scales, 24‑31 scales from the mid of the lower end of the ear opening to the mental scale. On both sides 4‑7 scales between supralabial and enlarged subocular scale; 29‑46 scales
around 5th whorl; 20‑23 tail whorls; 16‑20 scales beneath 4th left toe; 14‑20 preanofemoral pores
on either side.
Head covered with irregularly arranged scales of different size; the smallest situated above the
eyes. In the occipital region slightly pointed scales. Scales of the neck very small. Nostrils big. Anterior margins of the ear openings covered with enlarged triangular, pointed scales (2‑7 on either
side). Scales of the underside of the head small. Scales of the chest, the belly and the underside
of the extremities smooth. Scales of the soles slightly keeled. Subdigital scales with 3 or 4 distinct
keels. Scales of the back small and smooth. At the flanks enlarged lateral tubercles. Lateral sacral region with distinctly enlarged tubercular scales. Scales of the upper side of the upper arms
smooth, the scales of the forearm smooth or slightly pointed. Scales on the back of the upper arms
distinctly enlarged, some slightly keeled. Dorsal parts of the hind legs covered with enlarged tubercular scales, between which are very small and smooth scales. Scales on the upper side of the feet
enlarged and distinctly keeled.
C o l o u r : Uromastyx aegyptia aegyptia has the ability to undergo a physiological colour
change. At high temperatures the animals have a light brown to light grey colour, with a black
throat and small black dots on the neck. Some individuals have an entirely black to dark blue head.
At low temperatures the animals are dark grey to nearly black. Juveniles have characteristic transverse rows of yellow to orange ocelli on their back. The main colour of the body is greyish brown.
D i s t r i b u t i o n : See Fig. 3. The nominotypic subspecies inhabits northern Egypt east of
the River Nile, the Sinai Peninsula, Palestine and extreme north-western Saudi Arabia (Wadi Sawawin, Jebel as-Sinfa).
Uromastyx aegyptia microlepis Blanford, 1874
Plates 4‑5
Uromastix microlepis Blanford, 1874. — Proceedings of the Zoological Society of London (1874): 658.
Uromastyx aegyptius microlepis. — Mertens 1956; Jahreshefte des Vereins für vaterländische Naturkunde in Württemberg 111
(1): 93.
Uromastyx microlepis microlepis. — Arnold 1987 (ex errore); Proceedings of the Symposium on the Fauna and Zoogeography
of the Middle East; Beihefte zum TAVO A 28: 249.
T y p e s p e c i m e n : Lectotype, adult , Iraq, Basrah, without date, Capt. Phillips,
BMNH 1946.8.14.55, designated by Wilms & Böhme (2000 a).
D e s c r i p t i o n : S i z e : Large-growing taxon, maximum total length exceeding 700 mm.
on either side.
eyes. In the occipital region slightly pointed and convex scales. Scales of the neck very small. Nostrils big. Anterior margins of the ear openings covered with enlarged triangular, pointed scales (3‑9
on either side). Scales of the underside of the head small. Scales of the chest, the belly and the underside of the extremities smooth. Scales of the soles slightly keeled. Subdigital scales with 3‑4 dis-
443
Fig. 3: Distribution of Uromastyx aegyptia aegyptia and Uromastyx aegyptia microlepis (after Wilms & Böhme 2000 a). For the distribution of U. a. microlepis in the UAE see Fig. 4. — Localities: 1: Wadi Sawawin at the Jebel as-Sinfa region; 2: Wadi Araba (Palestine
and Jordan); 3: vicinity of Hazeva (Araba Valley, Palestine); 4: Sinai (Jebel el-Tik); 5: El Turkmania near Maghara; 6: 16 km W of
Feiran Oasis; 7: Suez, Jebel Suez; 8: between Suez and Ismailia; 9: Baltim; 10: 120 km S of Cairo, 35 km S of Cairo, Khanka N of
Cairo; 11: Wadi Hof (Helwan), Wadi Digla; 12: Kutamiya region, Wadi Iseili; 13: vicinity of Hurghada; 14: Jol, Hadramaut; 15: Bin
Khautar, Hadramaut; 16: Wadi Qitbit; 17: Jiddat al-Harasis; 18: 100 km from Muscat; 19: Abu Dhabi (al-Hamran, Bada Zaid);
20: Ras al-Barr, Bahrain; 21: al-Jubail, 70 km N of Bahrain; 22: Dhahran; 23: Harad; 24: 100 km N of Riyadh; 25: al-Quwayiyah;
26: Wadi Sirra; 27: Wadi Fatimah; 28: Jeddah; 29: Yanbu al-Bahr; 30: Madinah; 31: Anaiza; 32: as-Sa’ira; 33: ad-Dibdibah; 34: Wadi
al-Miyah; 35: between al-Gaisumah (= al-Qaysumah) and Turaif; 36: Kuwait; 37: Sirri Island; 38: Rudkhaneh-ye Shapur; 39: Fao;
40: 50 km W of Basrah, 60 km W of Basrah, Basrah; 41: W of Karbala; 42: Baghdad; 43: Kirkuk; 44: ar-Rutbah.
tinct keels. Scales of the back very small and smooth. No enlarged lateral tubercles; occasionally
specimens from the northern parts of the area can have slightly enlarged tubercles exclusively at
the rear parts of the body, but they are much smaller than tubercles in U. a. aegyptia. Scales of the
upper side of the upper arms smooth, those of the forearm smooth or slightly pointed. Scales on
the back of the upper arms distinctly enlarged, some slightly keeled. Dorsal parts of the hind legs
covered with enlarged tubercular scales, between which are very small and smooth scales. Scales on
the upper side of the feet enlarged and distinctly keeled.
444
C o l o u r : Uromastyx aegyptia microlepis has the ability to undergo a physiological colour
change. At high temperatures the animals are a light brown to yellow or greenish colour, with
a black throat and small black dots on the neck and dorsum. Some individuals have an entirely
black to dark blue head. At low temperatures the animals are dark grey, nearly black. Juveniles have
characteristic transverse rows of yellow to orange ocelli on their back. The main colouration of the
body is greyish brown.
D i s t r i b u t i o n : See Fig. 3 and (for details of the distribution in the UAE) Fig. 4. The
border between the ranges of the subspecies U. a. aegyptia and U. a. microlepis is obviously east of
Wadi Araba in Palestine and Jordan, and east of Wadi Sawawin in the Jebel as-Sinfa region of Saudi
Arabia.
Uromastyx aegyptia microlepis lives in the deserts and semi-deserts of Arabia (Saudi Arabia,
Yemen, Oman, United Arab Emirates, Qatar, Kuwait) and in Jordan, Syria, Iraq and coastal Iran.
Uromastyx leptieni Wilms & Böhme, 2000 – Leptien’s spiny-tailed lizard
Plates 6‑8
Uromastyx leptieni Wilms & Böhme, 2000. — Herpetozoa 13 (3/4): 142.
Ty p e s p e c i m e n : Holotype, adult , United Arab Emirates, Wadi Siji, VI.1983, R. Leptien, ZFMK 52398.
D e s c r i p t i o n : S i z e : Maximum length was given as 53 cm (SVL 28.3 cm, paratype
BMNH 85.II.7.4) by Wilms & Böhme (2000 a). The largest specimen measured in summer
2005 was a male of 67.5 cm total length (SVL 37.5 cm), weighing 1562 g. The largest female had
58.3 cm total length (SVL 33.7 cm), weighing 978 g.
on either side (in some specimens very small or absent).
eyes. Occipital scales and scales on the neck small and slightly pointed; on the neck, mostly on its
sides, intermixed with enlarged triangular, pointed scales. Anterior margins of the ear openings
covered with enlarged pointed scales (3‑7 on either side). Scales of the dorsal parts of the forelegs
slightly keeled. Scales of the ventral parts of the upper arm very small; on the forearm triangular
and keeled. Scales of the soles distinctly keeled; each subdigital scale with 3‑4 keels. Forelegs and
back without enlarged tubercles. Flank scalation with a row of enlarged tubercular scales from the
sacral region almost to the insertion of the forelegs. Dorsal scales are small and smooth, but some
specimens may have a series of enlarged plate-like scales from the sacral region to half the bodylength along the vertebral column. The diameter of these plates may reach three to four times the
diameter of surrounding scales.
C o l o u r : In preserved specimens the main colour is olive-beige with small dark dots. Neck,
head and throat are black. In some specimens the throat is marbled with black and orange. Ventral
part of the forelegs, chest and belly are marbled with grey. Ventral parts of the hind legs and first half
of the tail are grey. Colouration of juveniles is red-brown to dark brown with a dark brown to black
net-like pattern or with small irregularly arranged dark and light spots. Active specimens are pale
brown to almost entirely white, while cold specimens are grey with a dark grey to almost black head.
D i s t r i b u t i o n : See Fig. 4. Uromastyx leptieni is known from east of the Hajar al-Gharbi
mountains in northern Oman (vicinity of Muscat to Musandam Peninsula), and from the northeastern United Arab Emirates. The westernmost locality in the UAE is not Jebel Ali, as stated by
445
Plates 3‑8: Photographs of Uromastyx specimens (photos by T. Wilms, unless stated otherwise). 3: Uromastyx aegyptia aegyptia from
Safawi, Jordan (photo: D. Modry). 4‑5: Uromastyx aegyptia microlepis; 4: specimen from the vicinity of Abu Kamal, Syria (photo:
D. Modry); 5: extraordinary coloured specimen from Mahazat as-Sayd, Saudi Arabia. 6‑8: Uromastyx leptieni from the United Arab
Emirates; 6: adult specimen from Jebel Jayah (photo: E.N. Arnold); 7: juvenile from study site near Abu Dhabi International Airport;
8: subadult from same locality.
Wilms & Böhme (2000 a), but approximately at 24°27'N 54°38'E (study site near Abu Dhabi International Airport). In the UAE the nearest records of U. aegyptia microlepis are Shanayel, located
at the Hmeem road, and the above-mentioned study site. The distance between these two localities
is approximately 50 km. In Oman, the closest records of the two species are Muscat for U. leptieni
and a locality about 100 km inland from Muscat for U. a. microlepis.
446
Fig. 4: Distribution of Uromastyx aegyptia microlepis in the United Arab Emirates and of Uromastyx leptieni in the United Arab
Emirates and Oman (based on Baha El Din 1996, Wilms & Böhme 2000 a, Tourenq 2005, pers. obs.). It is assumed that populations west of the left dotted line belong to U. a. microlepis and those east of the right dotted line belong to U. leptieni; the populations between both lines (indicated by question marks) need further study to clarify their taxonomic status. — Localities: 1: Shanayel; 2: study site near Abu Dhabi International Airport; 3: al-Khazna; 4: Sweihan; 5: al-Ain; 6: Munayi; 7: vicinity of ar-Rustaq;
8: Muscat. — Identification: U. a. microlepis: specimens from near Jebel Dannah, from 23°45'N 53°33'E and from al-Hamran /
Bada Zaid (Abu Dhabi) examined (Wilms & Böhme 2000 a); specimens from Shanayel (locality 1) identified on the basis of photographs taken by C. Tourenq and I. Barcelo. Uromastyx leptieni: specimens from the numbered localities 2‑5 identified by pers. obs.
(June/July 2005); from Jebel Ali, Tawi Bil Khabis, Jebel Jayah and Wadi Siji examined (Wilms & Böhme 2000 a).
New information on the morphology and ecology of Uromastyx leptieni
The following paragraphs present observations and data collected on 82 live specimens of U. leptieni by Thomas Wilms and Brien Holmes at a site near Abu Dhabi International Airport (simply
referred to as “study site” in the following) in the period between 11 June 2005 and 16 July 2005.
N a t u r a l h i s t o r y n o t e s : Little is known about the natural history of Uromastyx leptieni.
Cunningham (2000, 2001 a, 2001 b) provided data on the ecology of a Uromastyx population
from the vicinity of al-Ain (“Dhub valley” at 23°25'N 55°35'E). Based on the photographs published by Cunningham (2000, 2001 a), we are inclined to assign these animals to U. leptieni. This
assumption is supported by the known distribution of U. leptieni in the UAE and the fact that
specimens of Uromastyx from Sweihan, al-Khazna and al-Ain (near Jebel Hafeet) belong without
doubt to U. leptieni (T. Wilms pers. obs.).
B u r r o w s p e c i f i c s : Our own observations show that most U. leptieni burrows enter the
soil in a straight line for up to 1 m, with a slope of about 30‑45°. Further down, the burrow in
most cases turns to the left or to the right at approximately a right angle and then descends in the
447
Fig. 5: Temperatures inside and outside of a Uromastyx leptieni burrow at the study site near Abu Dhabi International Airport between 20 June and 11 July 2005.
form of a spiral or in a zigzag line. Reported burrow length is more than 180 cm with a depth of
90‑120 cm (Cunningham 2001 a), which can be confirmed by our own observations.
Data loggers were used to measure air temperatures in the habitat (at a height of 1 m above
ground level) and soil temperatures at the surface outside the burrows (just in front of the entrance)
as well as inside (80 cm deep in the burrow) during a period between 20 June and 11 July 2005.
Air temperatures varied between a maximum of 51.8 °C and a minimum of 22.5 °C, while the soil
surface temperatures were between 55.4 °C and 22.1 °C. Within the burrow the temperature was
relatively stable, with a maximum of 40.1 °C and a minimum of 32.8 °C, thus providing ambient
temperatures to the lizards and allowing survival even when temperatures outside the shelters may
reach lethal values (Fig. 5). Apart from shelter against high temperatures, the burrows offer protection from drought. All dug-up burrows entered the moist soil layer, which could be important for
hydroregulation, as Uromastyx skin seems to have the ability to absorb water from moist surroundings (Kevork & Al-Uthman 1972).
Existing burrows are obviously reoccupied after the death of the former occupant. In the vicinity of some burrows, remains of large Uromastyx can be found, excavated by the successor. This
finding implies the high ecological value of the burrows, supposedly because of the difficulties to
dig into extraordinarily hard soil. The habitats exploited by Uromastyx leptieni for burrowing at the
study site consisted of aeolianite outcrops, gravel plains (partly sandy) and low hills. We observed
that more than one burrow was regularly used by a particular animal.
D a i l y a c t i v i t y : So far, only data concerning daily activity during summer are available.
According to Cunningham (2000), who provided data for May and June, the animals spend an
average of 81.2 % of the daylight hours underground. During our observation period, a total of
117 activity records have been taken (see Fig. 6). The diagram shows that U. leptieni exhibits a
bimodal activity rhythm during summer time with the highest activity before 11:30 and a resting
448
Fig. 6: Activity profile of a Uromastyx leptieni population at the study site near Abu Dhabi International Airport between 15 and
29 June 2005 in correlation with mean air temperatures.
period during the hottest time of the day. Afternoon activity is very limited and restricted to the
time after 15:30.
B o d y t e m p e r a t u r e s : During the observation period, body temperatures (cloacal temperatures) of four hand-caught Uromastyx leptieni were measured within less than one minute of capture (only animals caught without chasing were included). The cloacal temperature of these active
animals was between 35.8 °C and 42.2 °C (see Table 1).
To obtain information on the maximum cloacal temperature tolerated, measurements were
taken in 32 lizards caught in cage traps, which had been exposed to the sun. Cloacal temperature
ranged between 34.9 °C and 47.3 °C. Despite a cloacal temperature of 47.3 °C, which is close to
the lethal temperature in U. acanthinura (48‑50 °C according to Vernet et al. 1988), this specimen appeared healthy and alert.
D i e t : According to Cunningham (2000, 2001 a), Uromastyx leptieni feeds on 10 different plant
species at “Dhub valley”: Pennisetum divisium, Stipagrostis plumosa, Aerva javanica, Leptadenia pyrotechnica, Moltkiopsis ciliata, Heliotropium kotschyi, Haloxylon salicornicum, Citrullus colocynthis,
Taverniera cuneifolia and Polygala erioptera, of which grasses of the family Gramineae were preferred. This preference for grasses is confirmed by our observations, which suggest that U. leptieni
avoids feeding on the abundant Zygophyllum qatarense. A very limited amount of animal matter is
consumed. Obviously, in June and July feeding occurs almost exclusively before 09:00.
At the study site, faecal samples were collected and U. leptieni tracks followed to determine
food preferences. The longest measured track of a specimen collecting plant material was 387 m.
This specimen, a large female (total length 51.8 cm, weight 782 g), started at her burrow, grazed
on several grasses and bushes before returning to her burrow.
P o p u l a t i o n s t r u c t u r e a n d d e n s i t y : Within the sample of 82 specimens caught at
the study site and subsequently examined in detail, the sex ratio of males to females was 1 : 1.43
449
Table 1: Cloacal temperatures of four hand-caught Uromastyx leptieni at the time of capture, in comparison with soil and air temperatures.
Time of capture
09:25
10:15
11:00
11:05
Cloacal temperature
40.5 °C
42.2 °C
36.5 °C
35.8 °C
Soil temperature
41.1 °C
49.7 °C
[no record]
32.6 °C
Air temperature
39.3 °C
40.0 °C
[no record]
35.1 °C
Fig. 7: Scatter plot of total length against body weight of 82 individuals of Uromastyx leptieni from the study site near Abu Dhabi
International Airport. Males and females are indicated by the respective symbols, juveniles by circles.
with only 12 % of the animals being juveniles. Males generally grow larger than females (t‑test:
t = 4.57, α = 0.01) with a maximum total length of 67.5 cm (females: 58.3 cm). There is also a
significant difference in body weight between adult males and females (t‑test: t = 5.84, α = 0.01),
with adult males (total length larger than 40 cm) ranging between 384 g and 1572 g (mean:
999 g), while adult females (total length larger than 39 cm) range between 278 g and 983 g (mean:
599 g). Fig. 7 shows total length of juvenile, male and female Uromastyx leptieni plotted against
weight.
It is well known that Uromastyx females loose a considerable portion (approx. 30‑45 %) of
their body weight after oviposition (Zari 1999, Wilms 2001). Some of the females caught were
very skinny and emaciated, so that a recent oviposition could be supposed. This assumption was
supported by oviposition observed in some of the females caught at the study site, which were kept
at the Breeding Centre of Endangered Arabian Wildlife, Sharjah (D. Egan, pers. comm.). This observation indicates that reproduction of U. leptieni in the UAE takes place at least until July.
During the course of the translocation project, in total 213 specimens of U. leptieni were
caught, and this number is considered to be nearly the whole adult population of the area in
question. Fig. 8 gives an overview of the distribution of the Uromastyx population on the study
site. Three core areas with exceptionally high densities of Uromastyx have been identified within
450
Fig. 8: Satellite image of the study site near Abu Dhabi International Airport, with outlines of the three core areas and capture localities of Uromastyx leptieni superimposed.
the site, covering 160 ha (Area 1), 130 ha (Area 2) and 13 ha (Area 3). Despite intensive searching, only a few specimens have been found outside these areas. With 64 individuals of U. leptieni
caught in Area 1, the density was one animal per 2.5 ha, while in Area 2 (131 individuals) and
Area 3 (14 individuals) the density was one animal per approximately one hectare (0.99 and 0.93,
respectively).
Vegetation density in Area 2 was assessed by randomly placing six vegetation plots (each
100 m2). Vegetation cover in three plots was lower than 1 %; in the other three plots it ranged between 2 % and 3 %. Area 1 was severely degraded and had a lower vegetation cover than Area 2.
Multivariate analyses of the taxa of the Uromastyx aegyptia complex
Cluster analysis
Distance phenograms based on metric and meristic characters of individual adult specimens of
Uromastyx aegyptia and U. leptieni (41 males, 48 females; definition of variables V 1 to V 9 see
451
Fig. 9: Distance phenogram resulting from the cluster analysis of 41 adult males (left) and 48 adult females (right) of the Arabian
taxa of the Uromastyx aegyptia complex.
Table 2) were calculated, for males and females separately, using the average linkage method
(Fig. 9 ). The cophenetic correlation coefficient is 0.842 for males and 0.857 for females, while the
γ‑coefficient is 0.712 for males and 0.739 for females.
In males (left phenogram), one specimen from Egypt (number 1; BMNH 97.10.28.213,
Baltim Delta) is separated from all other specimens at the first major dichotomy. We consider
this animal an aberrant specimen because of its extraordinarily long tail (102.83 % of the SVL).
The second dichotomy separates two branches, of which one encompasses all but one individual
U. leptieni and the other branch the remaining U. aegyptia and the single U. leptieni (number 20).
In females (right phenogram), three U. leptieni specimens (numbers 17, 30 and 48) are
separated by the first dichotomy from all other specimens. All three specimens have very long
452
Table 2: Definition of variables based on metric and meristic characters. To eliminate allometric effects, proportions have been
calculated using metric values.
Variable
V 1
V 2
V 3
V 4
V 5
V 6
V 7
V 8
V 9
Definition
[snout-vent-length] × 100 / [tail length]
[head width] / [head length]
number of tail whorls
number of scales beneath the 4th toe on the left side
[maximum tail width at the 5th whorl] – [tail width between the 4th and 5th whorl] / 2
[tail length] / [maximum tail width at the 5th whorl]
number of gular scales
number of scales between gular and inguinal fold
number of scales around the 5th whorl
Table 3: Factor loading on the first three principal components from a correlation matrix of variables V 1 to V 9 for individual
males and females.
Variable
V 1
V 2
V 3
V 4
V 5
V 6
V 7
V 8
V 9
Eigen values
Accumulated percent of trace
Males
Females
Component 1 Component 2 Component 3 Component 1 Component 2 Component 3
–0.298
0.339
–0.155
–0.348
0.189
0.615
0.456
–0.229
–0.406
0.190
0.387
0.787
–0.282
–0.487
0.322
–0.365
0.460
–0.135
0.299
0.203
–0.357
–0.223
0.297
0.573
0.332
0.552
–0.451
–0.291
–0.424
0.450
0.398
0.261
–0.288
–0.233
0.386
–0.233
0.439
–0.509
0.136
–0.148
0.147
0.566
2.579
1.611
1.131
2.504
1.479
1.308
29.4 %
47.8 %
60.6 %
28.4 %
45.2 %
60.0 %
tails (101.79 %, 109.26 % and 112.30 % of the SVL, respectively). The second major dichotomy
separates roughly U. leptieni from U. aegyptia, with three individual U. aegyptia (numbers 3, 4 and
6) clustering together with the majority of U. leptieni and one individual U. leptieni (number 11)
clustering with the remaining U. aegyptia.
Because females of closely related Uromastyx species are generally more similar to one another
than males, it is not surprising that the cluster analysis of the datasets of individual females of the
three nominal taxa did not show such a high degree of congruence with the supposed taxonomic
arrangement as those of males did.
Principal component analysis (PCA)
A principal component analysis was run using the variables V 1 to V 9 for the same individual
males and females separately (for factor loading on the first three principal components see Table 3). In the projection of the first two principal components the male specimens (Fig. 10, above)
group together according to prior specimen classification, providing evidence that the operational
taxonomic units (OTUs) involved represent in fact the three taxa U. a. aegyptia, U. a. microlepis
and U. leptieni. The PCA run with the datasets of individual females (Fig. 10, below) did not
clearly separate the OTUs but has produced an area of broad overlapping. This result is in accordance with the result of the hierarchical cluster analysis, showing that females of different OTUs are
much more similar among each other than are males.
Fig. 10: Ordination of the individual
41 adult males (above)
and 48 adult females
(below) of the Arabian
taxa of the Uromastyx
aegyptia complex on the
first two principal components (Variables: V 1
to V 9, see Table 2).
453
454
This group consists of four species (U. benti Anderson, 1894, U. ocellata Lichtenstein, 1823,
U. ornata von Heyden, 1827 and U. yemenensis Wilms & Schmitz, 2007) inhabiting the African
and Arabian littoral regions of the Red Sea as well as the coastal areas of the Gulf of Aden in Yemen
and southern Oman (Wilms & Böhme 2000 c). One species, Uromastyx macfadyeni, that has been
previously assigned to this group due to morphological similarities, is most probably not related to
the other members of the Uromastyx ocellata group (Amer & Kumazawa 2005, Wilms & Schmitz
2007).
Three of the four species within the group, U. benti, U. ornata and U. yemenensis, are known
to occur in Arabia. The taxonomic history in this group is confusing, leading to difficulties in the
proper identification of the species in question. According to Wermuth (1967), all taxa deserve
specific rank, but Arnold (1986) treats the taxa ornata and philbyi as subspecies of Uromastyx ocellata, and Lanza (1988) regards macfadyeni likewise as a subspecies of U. ocellata. Wilms (1995)
follows this point of view and treats ornata, philbyi and macfadyeni as subspecies of U. ocellata.
According to Moody (1987), “ocellatus and ornatus [are] nearly indistinguishable” (see also Joger 1987), and Schätti & Gasperetti (1994) stated that they “consider benti, ornata and philbyi
to be conspecific with U. ocellata Lichtenstein”. Schätti & Desvoignes (1999) follow this concept
and regard U. benti, U. ornata, and U. philbyi as junior synonyms of U. ocellata. The examination
of 148 specimens of the Uromastyx ocellata group, including type material of all involved taxa, by
Wilms & Böhme (2000 c) revealed that in fact four species (Uromastyx ocellata, U. benti, U. macfadyeni and U. ornata as well as the recently discovered U. yemenensis) are involved and that they
can easily be distinguished. Of these species, U. macfadyeni is obviously not closely related to the
other taxa, because it forms a strongly supported clade basal to and clearly outside the rest of the
species of the Uromastyx ocellata group (Wilms & Schmitz 2007). For a detailed discussion of the
taxonomy and a key to the species of the Uromastyx ocellata group, see Wilms & Böhme (2000 c).
Uromastyx benti (Anderson, 1894) – Bent’s spiny-tailed lizard
Plate 9
Aporoscelis benti Anderson, 1894. — Annals and Magazine of Natural History (6) 14: 376.
Uromastix (Aporoscelis) benti. — Anderson 1896; A Contribution to the Herpetology of Arabia, with a Preliminary List of the
Reptiles and Batrachians of Egypt: 33, 63, 79, 85.
Uromastix simonyi Steindachner, 1899. — Anzeiger der Akademie der Wissenschaften in Wien, Mathematisch-Naturwissenschaftliche Klasse 36: 143.
Uromastix philbyi. — Haas & Battersby 1959; Copeia 1959: 202 (synonymy fide Arnold 1986).
Uromastyx thomasi. — Al-Badry & Al-Safadi 1982; Proceedings of the Egyptian Academy of Sciences 34: 66 (synonymy fide
Schätti 1989).
Uromastyx ocellata benti. — Schätti & Gasperetti 1994; Fauna of Saudi Arabia 14: 369.
Uromastyx ocellata. — Schätti & Desvoignes 1999; The Herpetofauna of Southern Yemen and the Sokotra Archipelago: 39.
Ty p e s p e c i m e n : Lectotype, adult , Yemen, Wadi Hadramaut, without date, J. Anderson, BMNH 1946.8.11.72, designated by Wilms & Böhme (2000 c).
D e s c r i p t i o n : S i z e : Medium-sized species, maximum total length 360 mm, maximum
SVL 196 mm (NHMW 16174).
S c a l a t i o n : 143‑187 scales around mid-body, 66‑86 scales between gular and inguinal fold,
23‑33 gular scales, 19‑27 scales from the mid of the lower end of the ear opening to the mental
scale. On both sides 4‑7 scales between supralabial and enlarged subocular scale; 28‑38 scales
around 5th whorl; 22‑26 tail whorls; 11‑15 scales beneath 4th left toe. No preanofemoral pores.
Head covered with irregularly arranged smooth scales of different size. Anterior margins of
the ear openings covered with enlarged scales (3‑6 on either side). No enlarged tubercular scales
455
Fig. 11: Distribution of Uromastyx benti (after Wilms & Schmitz 2007). — Localities: 1: Azzan; 2: al-Mukalla; 3: Ras Fartak;
4: vicinity of Mirbat.
on neck, forelegs or body. Body scales small and smooth. Thighs without enlarged tubercular
scales. Dorsal parts of the lower leg covered with very small scales, beneath the knee conical tubercular scales and/or enlarged keeled scales. Ventral parts of the lower leg covered with smooth
scales, resembling the scales of the soles in size and shape. Tail long and narrow, approximately
82.41 ± 5.48 % of SVL. Median scales of the upper side of the tail strongly keeled, dorsolateral
scales of the tail with well-developed spines.
C o l o u r : Males: In preserved specimens, ground colour of back, tail and hind legs yellowish
brown. Tail without distinct pattern, hind legs with a turquoise and orange colouration. Back with
a pattern consisting of dark brown lines and dots, as well as 7‑9 rows of ocelli (ivory-coloured with
dark brown edges). Dorsal side of the forelegs anthracite-coloured with orange and green colour
elements. Hands yellowish brown. Head orange or blue, marbled with dark brown and black. Underside of the head anthracite-coloured with some orange dots. Ventral parts of forelegs and chest
marbled with grey. Belly with narrow grey/anthracite cross-bands. — Females: much paler in colouration, with a yellowish brown ground colour and a pattern of small dark brown lines and dots.
D i s t r i b u t i o n : See Fig. 11. Uromastyx benti occurs in southern and south-eastern Yemen,
from the vicinity of Azzan eastwards to the Hadramaut Valley and along the coast of the Arabian
Sea. In the Sultanate of Oman this species is only known from the vicinity of Mirbat in southwestern Oman (Seufer et al. 1998, Wilms & Hulbert 2000).
Uromastyx ornata von Heyden, 1827 – Ornate spiny-tailed lizard
Uromastyx ornatus von Heyden, 1827. — Atlas zu der Reise im nördlichen Afrika von Eduard Rüppell. Abtheilung 1. Reptilien: 1.
Two subspecies are recognised: Uromastyx ornata ornata and Uromastyx ornata philbyi.
456
Plates 9‑14: Photographs of Uromastyx specimens (photos by T. Wilms, unless stated otherwise). 9: Uromastyx benti, male from
the vicinity of Mirbat, Oman. 10‑14: Uromastyx ornata ornata from Sinai Peninsula, Egypt; 10: male from Wadi Feiran (photo:
D. Modry); 11: adult male, blue colour phase; 12: adult male, red colour phase; 13: pregnant female; 14: hatchling.
Uromastyx ornata ornata von Heyden, 1827
Plates 10‑14
Uromastyx ornatus von Heyden, 1827. — Atlas zu der Reise im nördlichen Afrika von Eduard Rüppell. Abtheilung 1. Reptilien: 1.
Uromastix oronatus [sic]. — Farag & Banaja 1980 (typographic error); Bulletin of the Faculty of Science, King Abdulaziz
University 4: 12.
Uromastyx ocellatus ornatus. — Arnold 1986; Fauna of Saudi Arabia 8: 393.
Uromastyx ocellata ornata. — Lanza 1988; Biogeographia 14: 420.
457
T y p e s p e c i m e n : Holotype, female, Saudi Arabia, Mohila (= al-Muwaylih), 1828,
E. Rüppell, SMF 10403.
SVL 196 mm (BMNH 97.10.28.199).
around 5th whorl; 20‑23 tail whorls; 11‑15 scales beneath 4th left toe; 7‑14 preanofemoral pores on
either side.
Scalation homogenous, without enlarged tubercular scales on head, neck and back. Head covered with irregularly arranged scales of different size; the smallest situated above the eyes. In the
occipital region slightly pointed and slightly keeled scales. Scales of the neck very small. Anterior
margins of the ear openings covered with enlarged triangular, pointed scales (5‑7 on either side).
Scales of the underside of the head small and smooth. Scales of the chest, the belly and the underside of the extremities smooth. Scales of the soles slightly keeled. Subdigital scales with 3‑4 keels.
Scales of the back small and smooth. No tubercles on body and forearms. Scales of the forearm
smooth or slightly keeled. Dorsal parts of the hind legs covered with enlarged conical tubercular
scales. Between those scales very small and smooth scales. Scales on the upper side of the feet enlarged and smooth or slightly keeled.
C o l o u r : Uromastyx ornata is a very variable species. Colour of the males is green, blue or
red, with an irregularly reddish brown net-like pattern and yellow spots on the back. Sometimes
yellow cross-bands are present. Venter with dark pattern. Females are not as colourful as males.
They are light brown with dark brown spots and sometimes light yellow or light red spots. Belly
without pattern, yellowish or white.
Uromastyx ornata philbyi Parker, 1938
Uromastyx philbyi Parker, 1938. — Annals and Magazine of Natural History (11) 1: 484.
Uromastyx ocellatus philbyi. — Arnold 1986; Fauna of Saudi Arabia 8: 416.
Uromastyx ornatus philbyi. — Arnold 1987 (ex errore); Proceedings of the Symposium on the Fauna and Zoogeography of
the Middle East; Beihefte zum TAVO A 28: 252.
Uromastyx ocellata philbyi. — Schätti & Gasperetti 1994; Fauna of Saudi Arabia 14: 369.
Ty p e s p e c i m e n : Holotype, , Saudi Arabia, southern Hejaz between mountains and
Rub al-Khali, between Makkah and Shabwa, H.St.J.B. Philby, BMNH 1946.8.14.65 (former
number: BMNH 1938.2.1.1).
SVL 205 mm (MZUF 27906).
around 5th whorl; 17‑22 tail whorls; 11‑14 scales beneath 4th left toe; 7‑14 preanofemoral pores on
either side.
Scalation homogenous and similar to that of Uromastyx o. ornata. Anterior margins of the ear
openings covered with 5‑7 enlarged scales.
C o l o u r : Similar to that of Uromastyx o. ornata.
D i s t r i b u t i o n : See Fig. 12. Uromastyx o. ornata occurs on the Sinai Peninsula and in adjacent north-western Saudi Arabia, as well as at the extreme southern tip of Israel (area of Elat). Uro-
458
Fig. 12: Distribution of Uromastyx ornata ornata and Uromastyx ornata philbyi (after Wilms & Böhme 2000 c); sites with sympatric occurrence of both subspecies are shown. — Localities: 1: Sharm el-Sheikh; 2: Dahab; 3: Tor; 4: Mount Sinai, Wadi Nassen,
Kara’ah; 5: Wadi Feiran; 6: Elat; 7: Wadi Makfa; 8: Mohila (= al-Muwaylih); 9: Jebel as-Sinfa; 10: Jeddah, Burayman, Bazzah;
11: Makkah; 12: Taif; 13: Jebel Lababa (18°01'N 42°02'E); 14: southern Hejaz (between Makkah and Shabwa); 15: Ju’ Amlah
(approx. 26 km NW of Sa’dah).
mastyx o. philbyi occurs in the mountains of western Saudi Arabia, from the Jebel as-Sinfa region
to the southern Hejaz. Wilms & Böhme (2000 c) provide a discussion of the overlapping distribution of the two subspecies and point out that, on the basis of the available collections, a decision
whether philbyi deserves specific rank or not is impossible.
Plates 15‑22: Photographs of Uromastyx specimens (photos by T. Wilms, unless stated otherwise). 15‑16: Uromastyx yemenensis
yemenensis; 15: holotype in dorsal view; 16: same in ventral view. 17‑19: Uromastyx yemenensis shobraki; 17: holotype in dorsal
view; 18: same in ventral view; 19: male from northern Yemen. 20‑22: Uromastyx thomasi from Ras Hilf, Masirah Island, Oman;
20: male; 21: mating pair (photo: F. Hulbert); 22: hatchling.
459
460
Uromastyx yemenensis Wilms & Schmitz, 2007 – Southern Arabian spiny-tailed lizard
Uromastyx yemenensis Wilms & Schmitz, 2007. — Zootaxa 1394: 12.
Two subspecies are recognised: Uromastyx yemenensis yemenensis and Uromastyx yemenensis shobraki.
Uromastyx yemenensis yemenensis Wilms & Schmitz, 2007
Plates 15‑16
Uromastyx yemenensis yemenensis Wilms & Schmitz, 2007. — Zootaxa 1394: 12.
Ty p e s p e c i m e n : Holotype, adult , Republic of Yemen, Abyan Governorate, vicinity of
Lodar (= Lawdar), don. 1985, I. Haikal, ZFMK 47861.
D e s c r i p t i o n : S i z e : Medium-sized species, maximum total length 332 mm
(BMNH 1946.8.11.68).
around 5th whorl; 23‑26 tail whorls; 15‑18 scales beneath 4th left toe; no preanofemoral pores.
Anterior margins of the ear openings covered with 3‑6 enlarged scales. For a more detailed
description see Wilms & Schmitz (2007).
C o l o u r : Males: In preserved specimens, ground colour of the back, tail and hind legs yellowish brown. Tail without distinct pattern, hind legs with very small dark brown dots. Back with
a pattern consisting of dark brown lines and dots; five distinct cross-bands without or with very
few patterns on the back. Dorsal side of the forelegs anthracite-coloured. Hands yellowish brown.
Head yellowish brown, marbled with dark brown. Underside of the head anthracite-coloured
with some yellowish brown dots. Ventral parts of forelegs and chest marbled with grey. Belly with
narrow grey/anthracite cross-bands. — Females are much paler in colour, with a yellowish brown
ground colour with a pattern of small dark brown lines and dots. Five pale cross-bands on the
back. The ground colour of the ventral side is a light yellowish brown.
Uromastyx yemenensis shobraki Wilms & Schmitz, 2007
Plates 17‑19
Uromastyx yemenensis shobraki Wilms & Schmitz, 2007. — Zootaxa 1394: 16.
Ty p e s p e c i m e n : Holotype, adult , Yemen, Mafraq al-Mukha, 13.5 km W of Mafraq,
5‑6.IV.1988, B. Schätti, ZFMK 48681.
D e s c r i p t i o n : S i z e : Medium-sized species, maximum total length 393 mm
(MHNG 2538.47).
S c a l a t i o n : 163‑207 scales around mid body, 79‑97 scales between gular and inguinal fold,
around 5th whorl; 24-27 tail whorls; 15‑17 scales beneath 4th left toe; no preanofemoral pores.
Anterior margins of the ear openings covered with 3‑5 enlarged scales. For a more detailed
description see Wilms & Schmitz (2007).
C o l o u r : In preserved specimens, dorsal surface of head, body and hind limbs dark brown,
tail lighter. Light brown roundish dots (diameter 4 or 5 scales) are present on the dorsum, tending
to form transverse rows. In addition, irregular light brown dots are present on the whole dorsum.
Colour of the hands not different from the colour of the forearm. Head dark brown, with light
brown pattern. Ventral side of the specimen yellowish brown. Ventral side of head and chest marbled with anthracite and dark brown. There is an anthracite-coloured stripe-like pattern on the
sides of the belly.
D i s t r i b u t i o n : See Fig. 13. Uromastyx yemenensis is distributed in the south-western parts
of Yemen, with the range extending from the vicinity of Lawdar west to the Red Sea and from
461
Fig. 13: Distribution of Uromastyx yemenensis yemenensis and Uromastyx yemenensis shobraki. — Localities: 1: Oued (= Wadi) Zabid;
2: Mafraq al-Mukha; 3: Ta’izz; 4: Oued (= Wadi) Tiban; 5: Lahj; 6: 50 mi from Aden; 7: Zinjibar; 8: Lawdar.
Zabid south to the Gulf of Aden. The areas of the subspecies U. y. yemenensis and U. y. shobraki are
separated by a series of mountains and mountain ranges (example: Jebel Manar, Jebel Sawraq, Jebel
Sabir, Jebel Dawran) with heights up to 3,200 m (Wilms & Schmitz 2007).
Uromastyx princeps group
This group includes the Arabian species Uromastyx thomasi and the Somalian species U. princeps
(O’Shaughnessy, 1880). The shared derived character is the extraordinarily short and broad tail
with only 9‑14 annuli.
Uromastyx thomasi Parker, 1930 – Thomas’s spiny-tailed lizard
Plates 20‑22
Uromastix thomasi Parker, 1930. — Annals and Magazine of Natural History (10) 6: 595.
Ty p e s p e c i m e n : Holotype, , Oman, Rub al-Khali, Dhofar, Bu Ju’ay, without date,
B. Thomas, BMNH 1946.8.14.43 (former number: BMNH 1930.6.30.2).
D e s c r i p t i o n : S i z e : Small species, maximum total length approximately 24 cm, maximum SVL approximately 19 cm.
on either side.
Snout short. Head, neck and body without enlarged tubercular scales. Head covered with irregularly arranged scales of different size; the smallest situated above the eyes. Scales of the under-
462
Fig. 14: Distribution of Uromastyx thomasi (after Wilms 1998). — Localities: 1: between Bawi and Salalah; 2: Thumrait (= Thamarit); 3: Bu (= Bin) Ju’ay; 4: al-Ajaiz; 5: Ras Duqm, Jiddat al-Harasis; 6: Wadi Hauf (= Wadi Haluf ); 7: Masirah Island.
side of the head small and smooth. Scales of the chest, the belly and the underside of the extremities smooth. Scales of the soles distinctly keeled. Subdigital scales with 3 or 4 distinct keels. Upper
side of forelegs and thighs of hind legs covered with enlarged triangular, slightly keeled scales. Size
of the scales of the anterior margin of ear opening equals those of the cheek. Body scales small and
subimbricate. Scalation of the back not homogenous with respect to scale size. Smallest scales along
the vertebral column, biggest at the flanks. Tail distinctly depressed, short and disc-shaped.
C o l o u r : Yellowish green with a dark net-like pattern. A broad red stripe runs along the
vertebral column from the neck to the first half of the tail. In some specimens the head can be an
orange to red colour. Ventral side yellowish or white. Neck and sides of the head of the juveniles
striped (black and white). Colour of the upper side of the body black with six lighter transverse
bands, between which are yellowish to orange-coloured ocelli. Tail dorsally brown with some large
black spots. Belly and throat white. Ventral side of the tail white with black dots.
D i s t r i b u t i o n : See Fig. 14. Uromastyx thomasi lives in coastal Oman.
DISCUSSION
The following discussion of the taxonomy of the genus Uromastyx is based on the results of our
morphological studies (Wilms & Böhme 2000 a, 2000 b, 2000 c; Wilms 2001). Recently Amer &
Kumazawa (2005) found that a phylogeny of the genus Uromastyx, established on the basis of molecular genetics, is mostly consistent with our results, but they found a few discrepancies. In their
study, U. macfadyeni clustered with the species of the African U. acanthinura group, whereas morphologically it is grouped within the Afro-Arabian U. ocellata group. Another difference between
463
the two approaches is the placement of the U. aegyptia group within the cladogram. Morphologically, it is regarded as the sister taxon of the U. acanthinura group, while genetically it clusters with
the Arabian U. ocellata clade (without U. macfadyeni). Further studies, combining morphological
and genetic methods, are needed to establish reliable phylogenetic relationships within the genus.
As pointed out previously, the Uromastyx ocellata complex consists of six closely related taxa (ocellata, ornata, philbyi, benti, yemenensis and shobraki; but see Amer & Kumazawa 2005 for the status
of macfadyeni), distributed around the Red Sea as well as in the coastal area of the Gulf of Aden
(Djibouti, Somalia, Yemen and Oman), of which four deserve specific rank (U. ocellata, U. ornata,
U. benti and U. yemenensis). According to Wilms & Böhme (2000 c), U. philbyi should be treated
as a subspecies of U. ornata until new evidence becomes available.
To provide an overview on the taxonomic problems relating to the proper identification of the
taxa within the U. ocellata group, it seems to be necessary to discuss the identity of U. macfadyeni,
despite the fact that this species occurs outside the geographical scope of the present paper. According to Schätti (1989), one specimen from Djibouti (MHNG 2394.100) corresponds well
with the type specimens of U. macfadyeni from the Guban (northern Somalia). He states that: “Ein
Tier der Originalserie (BMNH 1937.12.5.125, Borama) besitzt einzelne, weit auseinanderliegende
Ocellen auf dem Rücken, beim Holotypus verschmelzen diese zu einem Netzmuster” [One animal
from the original series (BMNH 1937.12.5.125, Borama) has single, widely separated ocelli on
the back, in the holotype they merge to a net-like pattern]. The examination of those animals revealed that in fact BMNH 1937.12.5.125 (Borama) agreed very well with MHNG 2394.100, but
unfortunately is not part of the specimens examined by Parker (1932). The original description
of U. macfadyeni by Parker is based on only four specimens: the holotype BMNH 1946.8.14.54
(former number: BMNH 1925.4.3.1) from near Berbera, British Somaliland; paratype BMNH
1946.8.14.52‑53 (former number: BMNH 1931.8.1.141‑142) comprising 2 specimens from
Dayah Shabell, 24 miles SE of Berbera; and specimen BMNH 1905.10.30.31 from near Berbera
(today in the Museum of Comparative Zoology, Harvard University).
Even Parker (1942) himself contributed to the nomenclatural confusion by identifying
14 Uromastyx collected in the Guban (northern Somalia) between 10°00'N to 11°15'N and
42°40'E to 43°15'E as U. macfadyeni. He did not state any collection numbers, but the only specimens from the Guban in the Natural History Museum London are inventoried under the numbers
BMNH 1937.12.5.117‑130. With the exception of BMNH 1937.12.5.120 (since 1968 deposited
in the San Diego State Collection, USA) all of these specimens were examined, and all proved to
belong exclusively to U. ocellata.
Apart from the holotype and the two paratypes still in the Natural History Museum London, only one additional specimen belonging to the taxon U. macfadyeni could be found there
(BMNH 1956.1.6.55 from near Heis, 20 miles W of Mait, Somaliland). The synonymisation of
U. macfadyeni with U. ocellata by Schätti & Gasperetti (1994) is obviously based on misidentified U. ocellata from the Guban (Wilms & Böhme 2000 c). In addition to these findings, Amer &
Kumazawa (2005) and also Wilms & Schmitz (2007) showed that U. macfadyeni is genetically differentiated from all other species of the U. ocellata complex and should therefore not be recognised
as a member of this complex.
Further to the problem of discriminating U. macfadyeni from U. ocellata, some authors have
pointed out that “ocellatus and ornatus [are] nearly indistinguishable” (Moody 1987). Wilms &
464
Böhme (2000 c) have shown that in fact both species can be distinguished by two main characters: Firstly, the anterior margin of the ear is always covered by large, triangular scales in U. ornata
which are always absent in U. ocellata, and secondly U. ocellata has smaller body scales (U. ocellata:
189‑256 scales around mid-body; U. ornata: 149‑185 scales around mid-body). For a more detailed discussion of the differences between those two taxa, leading to the conclusion that both can
be separated at specific rank, see Wilms & Böhme (2000 c).
Another taxonomic problem arose when examining the relationship between U. ornata and the
taxon U. o. philbyi. The main character to distinguish U. o. ornata from U. o. philbyi is the relatively
broader tail in the latter (ratio of tail length to maximum tail width at the 5th whorl: U. o. ornata
3.61‑5.3; U. o. philbyi 3.03‑3.96). Wilms & Böhme (2000) found that the ranges of both phenotypes
broadly overlap in the mountains of western Saudi Arabia, based on six specimens of the narrow-tailed
phenotype (U. o. ornata-type) from inside the range of U. o. philbyi, namely: BMNH 1986.436 from
Jeddah; BMNH 1975.519 from 21°39'N 39°13'E and BMNH 1979.960 from 21°45'N 39°15'E,
both in the vicinity of Burayman; BMNH 1976.1748 from Wadi Fatimah; BMNH 1985.884 from
km 115 of the Makkah by-pass, 21°15'N 39°55'E; MHNG 2457.34 from Jebel Hababa (= Jebel
Lababa), 18°01'N 42°02'E. These localities are 800‑1300 km further south than the type locality of
U. o. ornata. According to Arnold (1986), the northernmost locality of U. o. philbyi (Jebel as-Sinfa,
27°57'N 35°47'E) is not far from the type locality of U. o. ornata (al-Muwaylih, 27°41'N 35°27'E).
At Burayman (21°40'N 39°10'E; BMNH 1964.296) and at Jebel Lababa (18°01'N 42°02'E;
MHNG 2457.33) both phenotypes are obviously sympatric, likewise in the region of Makkah and
Jeddah (U. o. philbyi: MHNG 2536.49, MNHP 4318, BMNH 1985.882, BMNH 1985.884,
BMNH 1986.434).
Based on the specimens available, a final decision on the taxonomic status of U. o. philbyi is
not yet possible. Three hypotheses are suggested:
– Both taxa (U. o. ornata and U. o. philbyi) are the extremes of an intraspecific cline: U. o. philbyi
must be synonymised with U. o. ornata.
– There are no intermediate specimens in the region in question (no gene flow between the two
sympatric types): U. o. philbyi must be lifted to specific rank.
– There is a secondary contact zone with gene flow between the two types: U. o. philbyi must be
treated as a subspecies of Uromastyx ornata.
As yet, none of these hypotheses can be excluded with certainty, so we suggest that until more
specimens and/or ecological data are available, U. o. philbyi should be treated as a subspecies of
U. ornata.
Wilms & Böhme (2000 a) provide an overview of the taxonomy of the taxa of the U. aegyptia group.
Three taxa are considered as valid at specific rank (Uromastyx aegyptia Forskål, 1775, Uromastyx
leptieni Wilms & Böhme, 2000 and Uromastyx occidentalis Mateo, Geniez, López-Jurado & Bons,
1998) and the taxon microlepis Blanford, 1874 is assigned as a subspecies to U. aegyptia. The validity of U. aegyptia and U. microlepis was under debate for a long time. The diagnostic features (sensu
Blanford 1874) to distinguish both taxa are the lack of enlarged tubercular scales on the flanks and
the presence of skin folds at the sides of the neck covered with tubercles in U. microlepis. The second
character has been found insufficient to distinguish both taxa properly (Wilms & Böhme 2000 a).
465
Schmidt (1939) had pointed out that it is sometimes difficult to distinguish between both
forms by the presence or absence of enlarged scales on the flanks, due to the variability of this
character in some populations. In the following decades U. microlepis was considered to be a valid
species by many authors, and Mertens (1956) suggested that U. microlepis could be a subspecies
of U. aegyptia. However, until the 1980s this viewpoint was not adopted by other authors. Moody
(1987) suggested U. microlepis to be a synonym of the nominotypic subspecies, because “the only
character used in earlier literature to diagnose U. aegyptia (spiny scales on the flanks) is variable
throughout the range of both species; and no other diagnostic characters have been discovered”.
Moody’s point of view was followed by Joger (1987) and Schätti & Gasperetti (1994). Arnold
(1980) supposed that U. aegyptia and U. microlepis might be conspecific, and in subsequent papers
(Arnold 1986, 1987) he treated U. microlepis as a subspecies of U. aegyptia. Leviton et al. (1992)
and Wilms (1995) adopted this viewpoint and pointed out that it would not be justified to synonymise U. microlepis with U. aegyptia before more information on the geographic variability of
these taxa was available. Anderson (1999) treated U. microlepis as a synonym of U. aegyptia.
According to Wilms & Böhme (2000 a), all specimens belonging to U. aegyptia have enlarged
tubercles scattered in their flank scalation, whereas most of the examined specimens of U. microlepis lack this feature. Some specimens of U. microlepis, especially from the northern parts of the
range (e.g. Kirkuk, Iraq), have very few slightly enlarged lateral tubercles. These never reach the
size of the tubercles found in U. aegyptia and they have a different distribution on the animals’
body. If present in U. microlepis, the lateral tubercles are much smaller than those in U. aegyptia
and restricted to the rear parts of the body just before the insertion of the hind legs, whereas in
U. aegyptia they are scattered in the whole flank scalation. Anderson (1999) did not find any tubercles on the flanks of the specimen he had examined from the Gulf coast of Arabia.
Furthermore, both taxa have different counts of ventral scales between the gular and inguinal
fold. The values are 126‑158 (mean: 142) for U. aegyptia and 149‑193 (mean: 171.8) for U. microlepis. Additionally, U. microlepis is more colourful than U. aegyptia, with a yellow or greenish
colour in warmed-up adult specimens.
Further studies are needed to show whether or not both taxa interbreed at the contact zone of
their ranges. This potential contact zone should be expected east of Wadi Araba (Jordan and Palestine) and east of Wadi Sawawin in the Jebel as-Sinfa region (Saudi Arabia). The specimens examined from both localities are without any doubt members of U. aegyptia, while specimens from east
of that region are exclusively U. microlepis. Because of the overall similarity of both taxa and their
parapatric distribution we think it is justified to treat U. microlepis as a subspecies of U. aegyptia, at
least until reproductive isolation mechanisms between both taxa are demonstrated.
Another confusing fact was the presence of another Uromastyx species from extreme southeastern Arabia. We believe that Moody’s (1987) conclusion to synonymise U. microlepis with
U. aegyptia was based on data obtained from specimens of U. leptieni which indeed have enlarged
lateral tubercles. However, specimens from surrounding areas do not have even slightly enlarged
lateral tubercles and therefore fit the description of U. microlepis. Uromastyx leptieni can be distinguished from U. aegyptia by the combination of the number of ventral and mid-body scales (see
Wilms & Böhme 2000 a: Fig. 2) and by the different colouration and pattern of the juveniles.
As has been shown in the present paper, it is possible to discriminate U. leptieni from U. aegyptia
using a multivariable approach of morphological datasets. Nevertheless, to further clarify the taxonomic situation a molecular approach should be applied.
The third species in this group is U. occidentalis, which is only known from two individuals.
The main diagnostic character for this species is the lack of preanal and femoral pores. Without
any doubt, U. leptieni is the sister taxon of U. aegyptia.
466
ACKNOWLEDGEMENTS
We would like to thank H.H. Prince Bandar bin Saud, Secretary General, National Commission for Wildlife Conservation and Development (NCWCD), Riyadh, Saudi Arabia; H.E. Prof.
Dr A.H. Abuzinada, former Secretary General, NCWCD; Dr M. Shobrak, NCWCD; Prof. Dr
I.A. Nader, NCWCD; Dr H. Tatwani, NCWCD; Mr A. bin Amer Al-Kiyumi, Director General
of Nature Reserves, Muscat, Oman and Mr S.M. Al-Saady, Director of Research, Muscat, Oman,
for their hospitality and help during the fieldwork of T. Wilms and for issuing the permits required. Special thanks are due to Brien Holmes, al-Ain, UAE and Felix Hulbert, Eltville, Germany
for their very pleasant companionship in the field.
The first author would also like to thank I. Barcelo, ERWDA, Abu Dhabi, UAE; Dr C. Drew,
Parsons International Limited, Abu Dhabi; Dr I. Galal, Jeddah, Saudi Arabia; Dr A.S. Gardner,
formerly Al-Koud, Muscat, Oman now Abu Dhabi, UAE; Dr B. Howarth, al-Ain, UAE; Mr
P. Paillat, formerly National Wildlife Research Center (NWRC), Taif, Saudi Arabia now National
Avian Research Center (NARC), Sweihan, Abu Dhabi, UAE; Dr C. Tourenq, ERWDA, Abu
Dhabi, UAE; P. Vercammen and D. Egan, Breeding Centre for Endangered Arabian Wildlife,
Sharjah, UAE and the teams of the NWRC, NARC and the King Khalid Wildlife Research Center, Thumamah, Saudi Arabia, for their hospitality and for the support provided.
We are grateful to the curators of the following museums for their hospitality and/or for the
loan of important collections: Dr E.N. Arnold, Dr C. McCarthy and Dr B. Clarke (BMNH); Dr
F. Tiedemann (NMW); Dr M. Poggesi and Dr C. Corti (MZUF); Dr V. Mahnert, Dr J. Mariaux
and Dr A. Schmitz (MHNG); Dr I. Ineich (MNHP); Prof. F.J. Obst and Dr U. Fritz (MTKD);
Dr K. Klemmer and Dr G. Köhler (SMF); Dr J. Hallermann (ZMH); Dr R. Günther (ZMB);
Dr U. Gruber and Dr F. Glaw (ZSM). Dr D. Modry (Brno, Czech Republic), Dr F. Krupp
(Frankfurt a.M., Germany) and Mr F. Hulbert (Eltville, Germany) and Dr E.N. Arnold (BMNH)
provided photographs.
Thomas Wilms is especially grateful to his brother, Matthias Wilms, who shared his knowledge on computers and mathematical science, and to the Wünstel family (Reptilium Landau) for
providing financial resources.
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Manuscript submitted:
Manuscript accepted:
03 April 2006
14 February 2007

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